Volatiles of the honey bee larva initiate oogenesis in the parasitic mite Varroa destructor

Summary. Varroa reproduction is closely synchronized to the development of its host. In this study we present a new bioassay for field and laboratory tests to evaluate host factors triggering Varroa oogenesis. Female mites deprived of feeding activated oogenesis when perceiving larval volatiles. In laboratory assays the living L5-larva and pentane extracts of the larval cuticle had a clear activating effect. Wax and larval food did not elicit Varroa oogenesis. The activating components apparently are in the polar fraction of the cuticular volatiles. The consequences of this regulative mechanism for the host parasite relationship and prospects for further research are discussed.     

Mating disruption of the honeybee mite Varroa destructor under laboratory and field conditions

Mating in the mite Varroa destructor, an ectoparasite of the honeybee, takes place within the sealed brood cell of its host and is triggered by a female sex pheromone consisting of three fatty acids and their respective ethyl esters. In a laboratory bioassay, we observed interactions among offspring of a Varroa female at 11 days after host cell capping in the absence and in the presence of the sex pheromone and clearly demonstrated that male mites were not able to distinguish between receptive daughters and either older or immature and unreceptive females when exposed to the pheromone. In addition, mating attempts in the presence of the pheromone with otherwise receptive females were clearly of shorter duration and males often failed to select a receptive female. In order to evaluate the effect of pheromone exposure on successful copulations and the number of spermatozoa transferred under field conditions, we sprayed one of the pheromone components, oleic acid, on an empty brood comb before host egg-laying and Varroa infestation and counted the spermatozoa of daughter mites taken from this comb. We could show that the number of spermatozoa was indeed reduced, and 20 % of females lacked spermatozoa. Our results open up new possibilities and represent a promising step toward biological control of Varroa mites in beehives.     

Female sex pheromone in immature insect males—a case of pre-emergence chemical mimicry?

Sexual selection by competition for mates is a formidable force that has led to extraordinary adaptations in males. Here we present results suggesting a novel case of pheromone mimicry in males of Lariophagus distinguendus, a parasitic wasp of beetle larvae that develop in stored grain. Females of L. distinguendus produce a pheromone even before they emerge from a grain. Males are attracted to the parasitised grain and wait for females to emerge. Males emerging later than others are under enormous selection pressure since females mate only once. We show evidence that developing males fool their earlier emerging competitors by mimicking the female pheromone. Males exposed to pupae of either sex exhibit typical courtship behaviour. Searching males are not only arrested by grains containing developing females but spend as much time on grains containing developing males. Hence, by distracting their competitors away from receptive females late males may increase their own chance to mate with these females. After emergence, males decompose the active compounds within 32 h probably to decrease molestation during their own search for mates. Chemical analyses of active pheromone extracts and bioassays using fractions demonstrate that the active compounds are among the cuticular hydrocarbons.     

Alkaloids and ascidian chemical defense: Evidence for the ecological role of natural products fromEudistoma olivaceum

The surface of the colonial ascidianEudistoma olivaceum (Van Name) is almost completely free of fouling organisms. I provide evidence that this epibiont-free surface is maintained by the alkaloids, Eudistomins G and H. These alkaloids were extracted from colonies collected in the shallow subtidal of Indian River Lagoon, Florida. USA, in November 1987. Concentrations of less than one-fifth of those found in the living ascidian inhibited the settlement of the larvae of five invertebrate species relative to appropriate controls in laboratory and field trials. Standard pharmacological assays have revealed that Eudistomins G and H were not the most biologically active of the eudistomins, indicating that these standard assays are not necessarily good predictors of antifouling activity. In an examination of possible alternate roles for these eudistomins, they proved ineffective as a fish antifeedant. PinfishLagadon rhomboides consumed agar pellets to which Eudistomins G and H had been added at concentrations 10- to 100-fold higher than those effective against settling larvae. These findings indicate that biologically active marine natural products may serve specific ecological roles.     

Sperm competition in the mite Caloglyphus berlesei

Summary The sperm utilisation pattern of C. berlesei was investigated using the sterile male method. The last male to mate with a female fertilised most of her eggs. In experiments where females mated with two virgin males, second males fertilised on average 86% of eggs when second matings took place immediately after the first. Last male sperm priority was also confirmed in experiments where females mated sequentially with three males. The proportion of eggs fertilised by the second male decreased to 71% and 56% when the interval between matings was 2 and 6 h, respectively. Males that had opportunities to mate prior to experiments copulated for a shorter time than virgin males, and second males fertilised on average 74% eggs.     

Choice in Varroa jacobsoni Oud. between honey bee drone or workerbrood cells for reproduction

Summary The effect of variation in copulation duration on sperm precedence (P₂) of the last male was investigated in a non-territorial libellulid using irradiated-male techniques and observations in a large field enclosure. Copulation duration varied between 6 min and more than 1 h (X = 23.19 min ± 12.74). In the clutch deposited immediately after the copulation, most copulation durations resulted in high sperm precedence for the last male. Several experimentally interrupted short matings still resulted in 100% precedence. However, most interrupted copulations of 10 min or less resulted in a low P₂. After long, uninterrupted copulations, P₂ remained high in clutches that were collected up to 9 days after the mating. After interrupted short copulations, P₂ fluctuated or remained at a low level from the first clutch onwards. This indicates that sperm mixing probably starts within 24 h and that after a long copulation virtually no rival sperm is left in the female's sperm storage organs. As most natural copulations are long, complete removal of rival sperm seems to be common in S. danae. Only 3 f 17 variables investigated explained a significant amount of the variation in copulation duration. Copulations shortened with increasing temperature and for each additional mating of the male. The strongest effect however, was due to time of day: early morning copulations lasted longer than midday copulations. Adjustment of the copulation duration to the decreasing chance of obtaining additional matings was found to be the most likely of five explanatory hypotheses that were evaluated.     

Population differences in male responses to chemical mating cues in the white-spotted longicorn beetle, Anoplophora malasiaca

Adult Anoplophora malasiaca beetles were collected from field populations inhabiting three different host plants: Citrus, Salix and Vaccinium. Male orientation responses towards odours of wounded branches of each host plant were examined. Males of all three populations were attracted most frequently to odours of their original hosts, indicating that male orientation behaviour was induced by the odours of the plants from which they were originally collected. We also observed the mating responses of the males to elytra extracts from young and mature females of the three populations and chemically analysed those extracts. Citrus males were highly responsive to female extracts, regardless of origin. Salix males were less reactive than Citrus males, but responded moderately to female extracts of all populations. Vaccinium males were never attracted to Citrus female extracts and responded more to extracts from mature Vaccinium and Salix females, indicating that males might distinguish female origin and maturity. All eight hydrocarbons, four ketones, and three lactones that were previously identified as female contact sex pheromones were found in the extracts of three populations and both ages. Higher amounts of lactones on mature females likely caused greater male responses. These results suggested that males would be more likely to approach a female feeding on the same host plant and subsequently recognise her sexual maturity based on the profile of contact sex pheromones on her elytra.     

Ontogenetic differences in chemical alarm cue production determine antipredator responses and learned predator recognition

How individuals assess, respond and subsequently learn from alarm cues is crucial to their survival and future fitness. Yet this information is not constant through time; many individuals are exposed to different predators throughout their life as they outgrow some predators or move to habitats containing different predators. To maximise overall fitness, individuals should discriminate between different cues and respond and learn from only those that are relevant to their current ontogenetic stage. We tested whether juvenile spiny chromis, Acanthochromis polyacanthus, could distinguish between chemical alarm cues from conspecific donors of different ontogenetic stages and whether the cue ontogenetic stage of the cue donor affected the efficacy of learning about predators. Juveniles displayed a significant antipredator response when conditioned with juvenile chemical alarm cues paired with predator odour but failed to respond when conditioned with predator odour paired with either adult alarm cues or with saltwater. Subsequently, individuals only recognised the predator odour alone as a threat when conditioned with juvenile alarm cues. This demonstrates that prey may be highly specific in how they use information from conspecific alarm cues, selectively responding to and learning from only those cues that are relevant to their developmental stage.     

Spectral tuning and perceptual differences do not explain the rejection of brood parasitic eggs by American robins (Turdus migratorius)

By laying their eggs in the nests of other birds, avian brood parasites impose the cost of rearing young upon their hosts. The recognition and rejection of foreign eggs are primary host defenses against costly brood parasitism. Hosts of parasitic brown-headed cowbirds (Molothrus ater) challenge coevolutionary theory because most cowbird hosts accept parasitic eggs despite their drastically different appearance from the hosts’ own eggs. American robins (Turdus migratorius) are one of only 10 % of the over 200 potential cowbird host species to robustly reject parasitic eggs, but the mechanisms driving the sensory bases of foreign egg rejection remain elusive. Our research combined avian visual perceptual modeling and behavioral experimentation to investigate chromatic cues eliciting parasitic egg rejection in American robins. We assessed the effects of perceivable background color differences between real host and model parasite eggs, across all four avian photoreceptors, on rates of rejection of model eggs spanning in color across the entire avian spectral sensitivity range, and including immaculate model eggs matching the natural colors of robin and cowbird eggs. The results suggest that egg rejection in robins is driven by the overall perceivable difference in color between own and artificial eggs, and input from all four single-cone avian photoreceptors affects the rejection decision. The results, however, also reveal that when viewed by the avian eye, natural cowbird eggs appear more similar in background color to robin eggs than predicted by the high rejection rate of these parasitic eggs. This suggests that robins respond specifically to parasitism by cowbirds, despite an apparent lack of sensory tuning toward the detection of the background color of cowbird eggs.     

The adaptive significance of male polymorphism in the acarid mite Caloglyphus berlesei

Summary Two forms of males occur in the acarid mite, Caloglyphus berlesei. One of them uses its thickened third pair of legs to kill other males; the other, without modified legs, does not attack other males. Previous studies have shown that the form of the male is environmentally determined: fighter males develop only at low population densities and their development can be suppressed by substances emanating from dense colonies. In this study, the duration of development, longevity and virility of the two male morphs were measured. The only significant difference was that fighters had a shorter development time between the two last moults. In another experiment, two groups of colonies were maintained under the same conditions but had different numbers of individuals: small colonies contained 2 fighters and 2 non-fighters, whereas large colonies contained 30 males of each type. The relative reproductive success of both morphs was estimated from numbers of matings. In small colonies, the estimated reproductive success (ERS) was significantly higher for fighters than for non-fighters, mainly because in over 50% of these colonies a single fighter male managed to kill all the rival males and monopolize the females. The opposite was true in large colonies, in which non-fighters achieved significantly higher ERS, mainly because the fighters were killed in fights more often than the non-fighters. This implies that the ratio of costs to benefits obtained by adopting the fighter-male strategy increases with the number of rival males, which may explain conditional male development in C. berlesei.     

Evidence for the remobilisation of transuranic elements in the terrestrial environment

The transuranium elements, Np, Pu and Am discharged from the BNFL fuel reprocessing plant at Sellafield have accumulated in the local environment. The processes responsible for their dispersal rely both on physical transportation and their chemical reactivity. The transuranics have a complex chemistry, with multiple oxidation states and a strongly polarising character. In the environment, the particle active III/IV and more mobile VNI oxidation state groups are important and govern their geochemical behaviour and subsequent dispersal. Studies of the behaviour of the transuranics, particularly Pu, in the Irish Sea, have shown that the majority of the radionuclides in the liquid effluent discharged from Sellafield, quickly becomes associated with the marine sediments. Their dispersal and distribution in the environment is then governed primarily by the movement of particulate material and for some sites it has been suggested that sediment profiles preserve the historical record of discharges from the plant. In tidally inundated soils, radionuclide levels are greatly enhanced. These soils are water-logged for long periods of the year, are strongly anoxic and accretion rate are very low. The distribution of Np, Pu and Am in the soil suggests that simple sedimentary accumulation mechanism cannot provide an adequate explanation for the profiles observed. From preliminary studies of soil pore water composition and detailed analysis of the variation of isotopic ratios in the soil cores, it is apparent that a small but significant component of the radionuclide inventory is mobile. In addition, it is clear that the mechanisms responsible for this mobility allows differentiation between the transuranium nuclides.     

Interpopulational differences in chemical composition and chemosensory recognition of femoral gland secretions of male lizards Podarcis hispanica: implications for sexual isolation in a species complex

Summary. Molecular studies suggest that the Iberian wall lizard, Podarcis hispanica, forms a species complex with several monophyletic types. In Central Spain two of these types are spatially not isolated and may interact. Sex pheromones are important for species recognition and, thus, differences between lizards’ types in chemicals used in intraspecific communication could lead to reproductive isolation. Analyses by GC-MS showed that the femoral gland secretions of adult males of different types were different. Males of one type had twelve exclusive compounds, and proportions of some shared compounds differed. This presumably would reflect selection for the persistency and efficiency of chemical signals in different environments; less volatile compounds and with a higher chemical stability being favoured in the type of lizards inhabiting more humid climatic conditions. Differential tongue-flick rates to scents from femoral secretions indicated that males were able to detect and discriminate between males of different types based on chemical cues alone. In contrast, females detected but did not seem able to discriminate between scents of the two types of males. Thus, multiple factors might be simultaneously acting either against or in favor of speciation, leading, on the one hand, to genetic differences between types, but, on the other hand, probably precluding an effective reproductive isolation in areas where both types of lizards may interact.     

Evidence for facultative heterotrophy in cultured zooxanthellae

The locus of symbiotic dinoflagellates within host cells provides a habitat which could potentially be exploited by the alga through heterotrophic uptake of host-derived organic substrates. Using zooxanthellae (Symbiodinium sp.) isolated from the tropical sea anemone Aiptasia pulchella collected from Kaneohe Bay, Hawaii, the effect of various potential organic substrates on growth in vitro was assessed in Erdschreiber seawater medium (ES) supplemented with organic compounds. Zooxanthellae maintained at 5 to 7 E m^-2 s^-1 (below compensation irradiance) grew heterotrophically when supplied with 100 M glycerol, glycolate, acetate, malate, or propionate, and grew in darkness on 100 M propionate. Zooxanthellae exposed to irradiance below compensation were able to utilize carbon sources in the unsupplemented ES medium for slow growth, but generally the growth rate of cultured zooxanthellae was a function of incubation irradiance. Zooxanthellae incubated for 10 wk in unsupplemented ES at 5 to 7 E m^-2 s^-1 were capable of growth at this low irradiance, but were also capable of net photosynthetic oxygen production at higher irradiances. This suggests that zooxanthellae can be photoautotrophic or facultatively heterotrophic. An estimate for the duration of mitosis (t _d ) is made on the basis of growth rate of cultured zooxanthellae in log-phase; this estimate of t _d =4.88 h is less than half the estimated t _d for zooxanthellae in situ.     

Delayed plumage maturation in Lazuli buntings: tests of the female mimicry and status signalling hypotheses

The evolutionary importance of delayed plumage maturation (DPM) in passerines, the condition when more than 1 year is required to achieve adult-like coloration, remains highly contentious. Adaptive hypotheses propose that aggression from after 2nd-year (ASY) males or predation favors DPM in 2nd-year (SY) males, thereby increasing SY male survivorship or reproductive success. However, each hypothesis suggests a distinct selective mechanism explaining “how” this is accomplished. Alternatively, DPM may be a consequence of a nonadaptive molt constraint. We tested the female mimicry and status signalling hypotheses in territorial ASY male lazuli buntings (Passerinaamoena) using three sets of model presentation experiments. The female mimicry hypothesis proposes that dull SY male plumage deceptively mimics female plumage, and predicts that ASY males can not distinguish SY male from female plumage. The status signalling hypothesis proposes that dull SY male plumage honestly signals low competitive threat, and predicts that ASY males respond less aggressively to dull versus bright, ASY-like plumage. Contrary to the female mimicry hypothesis, ASY males distinguished between SY male and female plumage, as they were aggressive to SY male models exclusively and attempted to copulate with female models. Supporting the status signalling hypothesis, ASY males were significantly less aggressive to SY versus ASY male plumage. While DPM may result from a physiological constraint on bright SY male plumage, our results support the idea that dull plumage in an SY male's first breeding season may be maintained by selection to reduce aggression from ASY males, serving as a signal of competitive status. Received: 21 February 1997 / Accepted after revision: 16 June 1997     

Genotypic variation in the parasitic dinoflagellate Hematodinium perezi along the Delmarva Peninsula,Virginia

Hematodinium perezi (genotype III) is a parasitic dinoflagellate that infects blue crabs along the eastern seaboard and Gulf of Mexico, USA. In order to examine the intra-specific genetic variation of this parasite, eleven microsatellite markers from H. perezi (III) were amplified from 227 infected blue crabs collected during 2008–2009 from six sites in Virginia. Simultaneous infections with multiple genetic types in a single-host individual were common and observed in 42 % of the samples. The remaining 58 % of samples had a single allele per locus at all eight polymorphic loci suggesting that the life history stages of the parasite in the host hemolymph are likely haploid. The composition and distribution of multi-locus genotypes (MLG) from samples with infections of a single genetic type indicated high genotypic variation along the Delmarva Peninsula, Virginia, with no evidence of population structure. The lack of linkage disequilibrium combined with the large number of unique MLGs (84 %) is strong evidence for recombination in the life cycle, but the sexual stages remain undetermined. This is the first evidence of ploidy level, infections by multiple genetic types in an individual host animal, high levels of genotypic variation, and sexual reproduction for any species of Hematodinium.     

Precise sex ratio in a parasitic wasp: the result of counting eggs

Summary The parasitic wasp Nesolynx albiclavus maintains a very skewed offspring sex ratio (ca. 4% males). We studied proximate, causal factors of the son-daughter choice mechanism to obtain insight into the precision of the procedures and the underlying rules. We analyzed the number of males in clutches produced during sessions of 12 or 24 h. The results show that the timing of the next male egg depends on the time elapsed and the number of eggs laid since the previous one. The influence of the passage of time per se during non-laying periods was reduced in an experiment with 4-h periods. The data show that the first male egg is produced by a fairly rigid process: about the sixth egg in a run will be left unfertilized. No further male egg will appear in the next 10–20 eggs if they are laid in rapid succession.     

Delayed maturation in plumage colour: Evidence for the female-mimicry hypothesis in the kestrel

Summary In many sexually dichromatic species, young males have female-like plumage during their first potential breeding year. The female-mimicry hypothesis (FMH) supposes that by possessing female-like plumage young males deceive older conspicuous males into believing that they are females, thus reducing competition from adult males. The status-signalling hypothesis (SSH) supposes that adult males can distinguish sex, but postulates that young males reduce competition from adult males by reliably signaling low status with their dull plumage. We tested these hypotheses in the European kestrel (Falco tinnunculus). Female-like young males settled to breed closer to adult males than did other adult males (Figs. 1a, b). By settling near adult males, young males seemed to increase their chance of mating with adult females. Adult female-young male pairs had better reproductive success than yearling-yearling pairs. These results suggest that there is an adaptive value in possessing a female-like plumage colour in the breeding season. To test the FMH, we measured sexual preference of adult males when adult females and young males were simultaneously shown in an aviary. Adult males were unable to recognize sex, because in half the cases they preferred young males (Fig. 3). However, when adult males and females were shown simultaneously, males preferred females (Fig. 2). Our results support the FMH rather than the SSH, because young males successfully deceived older males by their plumage.     

Modification of the activated sludge model for chemical dosage

Full-scale experiments have been carried out to adapt the activated sludge model ASM2d to include the influence of metal dosage (Fe³⁺ and Al³⁺) for phosphorus removal. Phosphorus removal rates, nitrification rates, as well as pH and sludge settling performance, were evaluated as functions of the metal dosages. Furthermore, models relating certain parameters to the dosage of chemicals have been derived. Corresponding parameters in the ASM2d and the secondary settler models, included in the Benchmark Simulation Model No 1 (BSM1), have been modified to take the metal influence into consideration. Based on the effluent limits and penalty policy of China, an equivalent evaluation method was derived for the total cost assessment. A large number of 300-day steady-state and 14-day open-loop dynamic simulations were performed to demonstrate the difference in behavior between the original and the modified BSM1. The results show that 1) both in low and high mole concentrations, Fe³⁺ addition results in a higher phosphorus removal rate than Al³⁺; 2) the sludge settling velocity will increase due to the metal addition; 3) the respiration rate of the activated sludge is decreased more by the dosage of Al³⁺ than Fe³⁺; 4) the inhibition of Al³⁺ on the nitrification rate is stronger than that of Fe³⁺; 5) the total operating cost will reach the minimum point for smaller dosages of Fe³⁺, but always increase with Al³⁺ addition.     

Evidence for the dominance of indirect effects in 50 trophic ecosystem networks

Indirect effects are powerful influences in ecosystems that may maintain species diversity and alter apparent relationships between species in surprising ways. Here, we applied network environ analysis to 50 empirically-based trophic ecosystem models to test the hypothesis that indirect flows dominate direct flows in ecosystem networks. Further, we used Monte Carlo based perturbations to investigate the robustness of these results to potential error in the underlying data. To explain our findings, we further investigated the importance of the microbial food web in recycling energy-matter using components of the Finn Cycling Index and analysis of environ centrality. We found that indirect flows dominate direct flows in 37/50 (74.0%) models. This increases to 31/35 (88.5%) models when we consider only models that have cycling structure and a representation of the microbial food web. The uncertainty analysis reveals that there is less error in the I/D values than the ±5% error introduced into the models, suggesting the results are robust to uncertainty. Our results show that the microbial food web mediates a substantial percentage of cycling in some systems (median = 30.2%), but its role is highly variable in these models, in agreement with the literature. Our results, combined with previous work, strongly suggest that indirect effects are dominant components of activity in ecosystems.