Plant secondary chemistry mediates the performance of a nutritional symbiont associated with a tree-killing herbivore

Many herbivores consume microbial food sources in addition to plant tissues for nutrition. Despite the ubiquity of herbivore-microbe feeding associations, few studies examine how host plant phenotypes affect microbial symbionts of herbivores. We tested the hypothesis that chemical polymorphism in a plant population mediates the performance of nutritional microbial symbionts. We surveyed the composition of ponderosa pine resin in northern Arizona, USA, for variation in six monoterpenes, and we approximated four chemical phenotypes. We reared populations of an herbivorous tree-killing beetle (Dendroctonus brevicomis) in ponderosa pine host material, controlling for three monoterpene compositions representing an alpha-pinene to delta-3-carene gradient. Beetles were reared in host material where the dominant monoterpene was alpha-pinene, delta-3-carene, or a phenotype that was intermediate between the two. We isolated nutritional fungal symbionts (Entomocorticium sp. B) from beetle populations reared in each phenotype and performed reciprocal growth experiments in media amended to represent four "average" monoterpene compositions. This allowed us to test the effects of natal host phenotype, chemical polymorphism, and the interaction between natal host phenotype and chemical polymorphism on a nutritional symbiont. Three important findings emerged: (1) fungal isolates grew 25-32% faster when acquired from beetles reared in the intermediate phenotype; (2) the mean growth rate of nutritional fungi varied up to 44% depending on which monoterpene composition media was amended with; and (3) fungal isolates uniformly performed best in the intermediate phenotype regardless of the chemical composition of their natal host. The performance of nutritional fungi related to both the chemical "history" of their associated herbivore and the chemical phenotypes they are exposed to. However, all fungal isolates appeared adapted to a common chemical phenotype. These experiments argue in favor of the hypothesis that chemical polymorphism in plant populations mediates growth of nutritional symbionts of herbivores. Intraspecific chemical polymorphism in plants contributes indirectly to the regulation of herbivore populations, and our experiments demonstrate that the ecological effects of plant secondary chemistry extend beyond the trophic scale of the herbivore-plant interaction.     

Learning and natal host influence host preference,handling time and sex allocation behaviour in a pupal parasitoid

The host choice and sex allocation decisions of a foraging female parasitoid will have an enormous influence on the life-history characteristics of her offspring. The pteromalid Pachycrepoideus vindemiae is a generalist idiobiont pupal parasitoid of many species of cyclorrhaphous Diptera. Wasps reared in Musca domestica were larger, had higher attack rates and greater male mating success than those reared in Drosophila melanogaster. In no-choice situations, na?ve female P. vindemiae took significantly less time to accept hosts conspecific with their natal host. Parasitoids that emerged from M. domestica pupae spent similar amounts of time ovipositing in both D. melanogaster and M. domestica. Those parasitoids that had emerged from D. melanogaster spent significantly longer attacking M. domestica pupae. The host choice behaviour of female P. vindemiae was influenced by an interaction between natal host and experience. Female P. vindemiae reared in M. domestica only showed a preference among hosts when allowed to gain experience attacking M. domestica, preferentially attacking that species. Similarly, female parasitoids reared on D. melanogaster only showed a preference among hosts when allowed to gain experience attacking D. melanogaster, again preferentially attacking that species. Wasp natal host also influenced sex allocation behaviour. While wasps from both hosts oviposited more females in the larger host, M. domestica, wasps that emerged from M. domestica had significantly more male-biased offspring sex ratios. These results indicate the importance of learning and natal host size in determining P. vindemiae attack rates, mating success, host preference and sex allocation behaviour, all critical components of parasitoid fitness. Electronic Publication     

Sociality reduces individual direct fitness in a communally breeding rodent,the colonial tuco-tuco (<Emphasis Type="Italic">Ctenomys sociabilis</Emphasis>)

In many social vertebrates, remaining in the natal group leads to at least short-term reductions in the direct fitness of philopatric animals. Among communally breeding rodents, the direct fitness costs of philopatry appear to increase as the frequency of successful natal dispersal decreases, suggesting a functional link between constraints on natal dispersal and the reproductive consequences of sociality. To explore this relationship empirically, I documented patterns of direct fitness among female colonial tuco-tucos (Ctenomys sociabilis), which are group-living subterranean rodents from southwestern Argentina. Demographic data suggest that successful natal dispersal is rare in this species, leading to the prediction that natal philopatry in C. sociabilis is associated with significant reductions in individual direct fitness. Using data obtained during 1996–2001, I compared the direct fitness of females that dispersed from their natal group and bred alone as yearlings to that of females that lived and bred in their natal group as yearlings. Philopatric yearlings reared significantly fewer young to weaning than did disperser (lone) yearlings. Although neither survival to a second breeding season nor the estimated lifetime number of pups reared to weaning differed between dispersal strategies, the annual direct fitness of group-living females was 23–40% less than expected, suggesting that philopatric animals experienced a substantial direct fitness cost by remaining in their natal group. These data yield important insights into the adaptive bases for group living in C. sociabilis and suggest that constraints on natal dispersal are an important factor favoring group living in this species.Communicated by J. Wilkinson     

Consequences of communal gall occupation and a test for kin discrimination in the aphid Tamalia coweni (Cockerell) (Homoptera: Aphididae)

This study investigated the consequences of communal gall induction on individual and group fitness in the aphid Tamaliacoweni. The possibilities that kin discrimination and foundress density are factors favoring communal gall occupation were examined. Clonally produced aphid foundresses were collected to create two treatments: clonal groups and groups of less closely related individuals. These were confined on suitable host plant tissue to compare their respective propensities towards communal behavior. There were no significant differences in the frequencies of communal gall occupation; therefore, active kin discrimination by T.coweni foundresses apparently does not play a role in their communal behavior, within the context of this experiment. In a second experiment, aphid foundress density on the host plant was manipulated in three treatments and was correlated with the frequency of communal gall occupation. Individual fitness was inversely related to the mean number of foundresses per gall. These results suggest that communal gall occupation does not necessarily represent mutual cooperation but may instead be the outcome of competition for limited gall sites on the host plant. However, natural selection at the clonal level may favor communal gall occupation under the conditions of resource limitation and high within-group relatedness. Received: 26 September 1997 / Accepted after revision: 28 March 1998     

Fitness related diet-mixing by intraspecific host-plant-switching of specialist insect herbivores

Generalist insect herbivores may profit by feeding on a mixture of plant species that differ in nutritional quality. Herbivore performance can also be affected by intraspecific host plant variation. However, it is unknown whether conspecific plant individuals differ sufficiently to promote diet-mixing behavior in specialist herbivores. We experimentally tested this "specialist diet-mixing hypothesis" for specialist caterpillars (Chrysopsyche imparilis, Lasiocampidae) in a West African savanna. The caterpillars switched regularly between host tree individuals (Combretum fragrans, Combretaceae). To examine whether switching benefited caterpillar performance via diet-mixing, the caterpillars were reared either on leaves from several plant individuals (mixed diet) or on leaves from a single plant. The strongest effect of diet-mixing was found for fecundity, with females reared on a mixed diet laying significantly more eggs than sisters receiving a single-plant diet. In addition, a mixed diet decreased variability in egg size and increased the growth of second-instar caterpillars. Supplementary food choice experiments were conducted to assess a potential influence of lowered host quality (induced by herbivory) on caterpillar behavior; no such effect was found. By linking intraspecific host-switching behavior and herbivore performance, this study provides new information on the relevance of intraspecific plant variation for herbivorous insects.     

Ecological costs on local adaptation of an insect herbivore imposed by host plants and enemies

Herbivore populations may become adapted to the defenses of their local hosts, but the traits that maximize host exploitation may also carry ecological costs. We investigated the patterns and costs of local adaptation in the pine processionary moth, Thaumetopoea pityocampa, to its host plants, Pinus nigra and P. sylvestris. The two hosts differ in needle toughness, a major feeding impediment for leaf-eating insects. We observed a west-to-east gradient of increasing progeny size in the Italian Alps, matching the pattern in toughness of their respective local host plant. Eastern populations that feed on the native P. nigra with tough needles had larger eggs, and neonate larvae with larger head capsules, than western populations that feed on the native P. sylvestris and the introduced P. nigra with softer foliage. In a reciprocal transfer experiment that involved the eastern-most and the western-most populations of T. pityocampa from this region, and excluded natural enemies, we found evidence for local adaptation to the host plant. Specifically, larvae from the western population only performed well when raised on their local hosts with soft needles, and they suffered near-complete mortality on the tough foliage at the eastern site. In contrast, larvae from the eastern population survived equally well at both sites. Local adaptation involved a trade-off between progeny size and the number of offspring. We hypothesized that an additional cost, imposed by natural enemies, may be associated with increased egg size: we also observed a west-to-east gradient of increased egg parasitism. We tested this hypothesis in a common garden by exposing eggs of both populations to parasitism by two native egg parasitoids, Ooencyrtus pityocampae and Baryscapus servadeii. The eastern population suffered a higher level of parasitoid attack by O. pityocampae than the western population, and performance of hatched adults of both parasitoids was enhanced in large eggs. Thus, increased neonate quality (larger eggs yielding larger larvae) confers an advantage on tough foliage but incurs the ecological cost of increased parasitism, which may constrain further adaptation by this herbivore.     

Impact of an invasive oak gall wasp on a native butterfly: a test of plant-mediated competition

Phytophagous insects commonly interact through shared host plants. These interactions, however, do not occur in accordance with traditional paradigms of competition, and competition in phytophagous insects is still being defined. It remains unclear, for example, if particular guilds of insects are superior competitors or important players in structuring insect communities. Gall-forming insects are likely candidates for such superior competitors because of their ability to manipulate host plants, but their role as competitors is understudied. We investigate the effect of invasive populations of an oak gall wasp, Neuroterus saltatorius, on a native specialist butterfly, Erynnis propertius, as mediated by their shared host plant, Quercus garryana. This gall wasp occurs at high densities in its introduced range, where we stocked enclosures with caterpillars on trees that varied in gall wasp density. Biomass production of butterflies was lower in enclosures on high-density than on low-density trees because overwintering caterpillars were smaller, and fewer of them eclosed into adults the following spring. To see if the gall wasp induced changes in foliar quality, we measured host plant quality before and after gall induction on 30 trees each at two sites. We found a positive relationship between gall wasp density and the percentage change in foliar C:N, a negative relationship between gall wasp density and the percentage change in foliar water at one site, and no relationship between the percentage change in protein-binding capacity (i.e., phenolics) and gall-wasp density. Additionally, there was a negative relationship between foliar quality and butterfly performance. Our results provide evidence for a plant-mediated impact of an invasive oak gall wasp on a native butterfly and suggest that gall wasps could act as superior competitors, especially when they occur at high densities.     

Bottom-up and top-down effects on insect herbivores do not vary among sites of different salinity

It has been suggested, but rarely tested, that the relative strength of top-down and bottom-up factors in communities varies along an environmental stress gradient. We compared the strength of bottom-up and top-down effects on the densities of insect herbivores along a range of sites of different salinities in west-central Florida. We used a 2 x 2 factorial design with plots divided into four treatments: (1) bottom-up manipulation, where fertilizer was applied to increase plant quality; (2) top-down manipulation, where sticky traps were used to reduce the effects of natural enemies (parasitoids); (3) bottom-up and top-down manipulation, where fertilizer was applied and sticky traps were used; and (4) control plots. These plots were established along a range of salinities among seven different sites containing the salt marsh plant Borrichia frutescens. In each plot, we determined the parasitism levels and abundances of the sap sucker Pissonotus quadripustulatus, the gall maker Asphondylia borrichiae, and the lepidopteran stem borer Argyresthia spp. Gall density, Pissonotus density, and stem borer density were significantly higher in lower salinity sites, suggesting a strong effect of environmental stress. There was a significant increase of galls and Pissonotus and a marginally significant increase of bored stems on fertilized plots but not on trapped plots. There was a significant interaction of site and fertilizer on gall parasitism. There were no interactions of either treatment with salinity on herbivore densities. The general lack of interaction between salinity level and other treatments on herbivore densities contrasts with our previous result where treatment effects did vary with salinity level on a large experimentally generated salinity gradient at one site. Thus, the results of the present paper suggest that, while environmental stress can modify top-down and bottom-up effects on herbivores at single sites, variation in site-to-site factors, possibly including clonal identity of plant, affects herbivore densities so much as to swamp out any observable interaction between environmental stress and top-down or bottom-up factors.     

Condition-dependent sex allocation by clones of a galling aphid

Local mate competition (LMC) has been postulated to be the primary factor of female-biased sex allocation. In animals such as aphids that exhibit seasonal alternations of clonal and sexual reproduction, there is a high possibility of intra-clonal mating and LMC. This possibility is more plausible for more fecund clones, but outbreeding is predicted for less fecund clones. We hypothesize that clones that are more fecund will gain higher fitness returns by reducing investment in males because of more intense LMC among clonal males. We tested this hypothesis by elucidating the clonal sex allocation patterns of the galling aphid Kaltenbachiella japonica, in which inbreeding and LMC appear to be common. Winged mothers that emerge from a gall, belonging to the same clone, produced males and sexual females asexually on a branch, without dispersing to other trees. The heavier the gall, the more winged mothers were produced from the gall. Individual mothers produced a constant number of males and a variable number of females. The clonal sex allocation to males was 39.8?%, on average, and decreased with increasing gall weight. This result showed that clones that were more fecund exhibited more female-biased sex allocation and thus supported our hypothesis. Furthermore, our results corroborated Stubblefield and Seger’s hypothesis for sex allocation in patch structure rather than Yamaguchi’s constant male hypothesis. We conclude that K. japonica clones are able to adjust their sex allocation patterns adaptively depending on the quality of resources in the galls.     

Variability in host plant chemistry: behavioural responses and life-history parameters of the Colorado potato beetle (Leptinotarsa decemlineata)

Summary. Many studies investigating effects of plant chemicals on herbivore performance have reported contradictory results, perhaps because of possible interaction between different chemicals. Also, a herbivore’s performance is not necessarily consistent with its food or oviposition preference. Our aim was to investigate simultaneously antibiosis (larval growth and survival) and antixenosis (oviposition and feeding preferences) responses in herbivore to three plant chemicals, of which one is expected to have positive and two are expected to have negative effects. Antibiosis was measured by correlating the nitrogen and glycoalkaloid levels in host plants to the survival and adult size of Leptinotarsa decemlineata, by rearing larvae on whole plants of three potato varieties. Although host plants differed in their glycoalkaloid levels, survival rate and adult body size did not differ among beetles reared on different potato varieties. This suggests that beetles are quite robust for differences in both foliar α-chaconine and foliar α-solanine content. However, differences in antixenosis were found although they could not be directly predicted from the leaf chemistry. Females preferred to lay their eggs on the variety with high α-solanine content (Nevsky) towards which males showed a tendency to feeding preference. Overall, our results confirm that beetles are well adapted to the chemical defences of potato plants as potato varieties did not significantly affect beetle performance, but differences in oviposition preference may still result in major differences in the amount of damage inflicted on plants in the fields.     

Host-plant genotypic diversity mediates the distribution of an ecosystem engineer

Ecosystem engineers affect ecological communities by physically modifying the environment. Understanding the factors determining the distribution of engineers offers a powerful predictive tool for community ecology. In this study, we examine whether the goldenrod bunch gall midge (Rhopalomyia solidaginis) functions as an ecosystem engineer in an old-field ecosystem by altering the composition of arthropod species associated with a dominant host plant, Solidago altissima. We also examine the suite of factors that could affect the distribution and abundance of this ecosystem engineer. The presence of bunch galls increased species richness and altered the structure of associated arthropod communities. The best predictors of gall abundance were host-plant genotype and plot-level genotypic diversity. We found positive, nonadditive effects of genotypic diversity on gall abundance. Our results indicate that incorporating a genetic component in studies of ecosystem engineers can help predict their distribution and abundance, and ultimately their effects on biodiversity.     

Experimental evidence for effective and altruistic colony defence against natural predators by soldiers of the gall-forming aphid Pemphigus spyrothecae (Hemiptera : Pemphigidae)

Summary The thick-legged first instar soldiers of the gall-forming aphid Pemphigus spyrothecae Pass. are able to protect the aphids in the gall from being eaten by a range of insect predators. In artificial galls, the soldier aphids were able to kill first instar ladybirds Adalia bipunctata (L.) (Coleoptera: Coccinellidae), early third instar hoverfly larvae Eupeodes (Metasyrphus) corollae (Fab.) (Diptera: Syrphidae), and first-third instar Anthocoris nemoralis (Fab.) (Hemiptera: Cimicidae). Almost all the aphids that attacked the predators were themselves killed. The soldiers were also able to kill predators introduced into natural galls. Experiments were devised in which individuals of Anthocoris minki, which is the most important insect predator of the gall generations of Pemphigus at the study site, were free to enter and leave the gall: the soldiers were effective both in preventing the predator's access to the gall and in killing those predators that did manage to get in. In galls with experimentally manipulated numbers of soldiers and non-soldiers, it was clearly shown that it is the soldiers alone that kill the predators (Anthocoris minki and 1st instar Adalia bipunctata) (see Table 3). Even though many soldiers may die during these encounters, the selective advantage of killing the predators is high, since observations show that individual A. minki can pass through more than one instar inside a gall and kill all the aphids therein. The aphids were not observed to attack conspecific aphids from other galls or the cohabiting aphid Chaitophorus leucomelas Koch. The primary role of the soldier caste is therefore the defence of the aphid colony against predators.     

Genetic variation in defense chemistry in wild cabbages affects herbivores and their endoparasitoids

Populations of wild Brassica oleracea L. grow naturally along the Atlantic coastlines of the United Kingdom and France. Over a very small spatial scale (i.e., <15 km) these populations differ in the expression of the defensive compounds, glucosinolates (GS). Thus far, very few studies have examined interactions between genetically distinct populations of a wild plant species and associated consumers in a multitrophic framework. Here, we compared the development of a specialist (Pieris rapae) and a generalist (Mamestra brassicae) insect herbivore and their endoparasitoids (Cotesia rubecula and Microplitis mediator, respectively) on three wild populations and one cultivar of B. oleracea under controlled greenhouse conditions. Herbivore performance was differentially affected by the plant population on which they were reared. Plant population influenced only development time and pupal mass in P. rapae, whereas plant population also had a dramatic effect on survival of M. brassicae. Prolonged development time in P. rapae corresponded with high levels of the indole GS, neoglucobrassicin, whereas reduced survival in M. brassicae coincided with high levels of the aliphatic GS, gluconapin and sinigrin. The difference between the two species can be explained by the fact that the specialist P. rapae is adapted to feed on plants containing GS and has evolved an effective detoxification system against aliphatic GS. The different B. oleracea populations also affected development of the endoparasitoids. Differences in food-plant quality for the hosts were reflected in adult size in C. rubecula and survival in M. mediator, and further showed that parasitoid performance is also affected by herbivore diet.     

Variation in herbivore-mediated indirect effects of an invasive plant on a native plant

Theory predicts that damage by a shared herbivore to a secondary host plant species may either be higher or lower in the vicinity of a preferred host plant species. To evaluate the importance of ecological factors, such as host plant proximity and density, in determining the direction and strength of such herbivore-mediated indirect effects, we quantified oviposition by the exotic weevil Rhinocyllus conicus on the native wavyleaf thistle Cirsium undulatum in midgrass prairie on loam soils in the upper Great Plains, USA. Over three years (2001-2003), the number of eggs laid by R. conicus on C. undulatum always decreased significantly with distance (0-220 m) from a musk thistle (Carduus nutans L.) patch. Neither the level of R. conicus oviposition on C. undulatum nor the strength of the distance effect was predicted by local musk thistle patch density or by local C. undulatum density (<5 m). The results suggest that high R. conicus egg loads on C. undulatum near musk thistle resulted from the native thistle's co-occurrence with the coevolved preferred exotic host plant and not from the weevil's response to local host plant density. Mean egg loads on C. undulatum also were greater at sites with higher R. conicus densities. We conclude that both preferred-plant proximity and shared herbivore density strongly affected the herbivore-mediated indirect interaction, suggesting that such interactions are important pathways by which invasive exotic weeds can indirectly impact native plants.     

Photosynthesis and sink activity of wasp-induced galls in Acacia pycnantha

Although insect galls are widely known to influence source-sink relationships in plants, the relationship between photosynthesis and gall activity has not been extensively studied. In this study we used 14CO2, photosynthesis, and respiration measurements to examine the capacity of bud galls induced by the wasp Trichilogaster signiventris (Pteromalidae) as carbon sinks in Acacia pycnantha. Galls of this species develop either in vegetative or reproductive buds, depending on the availability of tissues at different times of the year, and effectively eliminate seed production by the plant. Photosynthetic rates in phyllodes subtending clusters of galls were greater than rates in control phyllodes, a result we attributed to photosynthesis compensating for increased carbon demand by the galls. Contrary to previous studies, we found that photosynthesis within galls contributed substantially to the carbon budgets of the galls, particularly in large, mature galls, which exhibited lower specific respiration rates allowing for a net carbon gain in the light. To determine the sink capacity and competitive potential of galls, we measured the proportion of specific radioactivity in galls originating from either vegetative or reproductive buds and found no difference between them. The proportion of the total amount of phyllode-derived 14C accumulated in both clustered and solitary galls was less than that in fruits. Galls and fruits were predominantly reliant on subtending rather than on distant phyllodes for photosynthate. Solitary galls that developed in vegetative buds constituted considerably stronger sinks than galls in clusters on inflorescences where there was competition between galls or fruits for resources from the subtending phyllode. Wasps developing in solitary vegetative galls were correspondingly significantly larger than those from clustered galls. We conclude that, in the absence of inflorescence buds during summer and fall, the ability of the wasps to cause gall formation in vegetative tissues tempers intraspecific competition and substantially increases the availability of plant resources for the development of wasps in such galls.     

Variation in composition of predator-attracting allelochemicals emitted by herbivore-infested plants: Relative influence of plant and herbivore

Summary During foraging, natural enemies of herbivores may employ volatile allelochemicals that originate from an interaction of the herbivore and its host plant. The composition of allelochemical blends emitted by herbivore-infested plants is known to be affected by both the herbivore and the plant. Our chemical data add new evidence to the recent notion that the plants are more important than the herbivore in affecting the composition of the volatile blends. Blends emitted by apple leaves infested with spider mites of 2 different species,T. urticae andP. ulmi, differed less in composition (principally quantitative differences for some compounds) than blends emitted by leaves of two apple cultivars infested by the same spider-mite species,T. urticae (many quantitative and a few qualitative differences). Comparison between three plant species — apple, cucumber and Lima bean — reveals even larger differences between volatile blends emitted upon spider-mite damage (many quantitative differences and several qualitative differences).     

Modelling the spatial population dynamics of the green oak leaf roller (Tortrix viridana) using density dependent competitive interactions: Effects of herbivore mortality and varying host-plant quality

Cyclic population dynamics of forest insects with periods of more than two generations have been discussed in relation to a variety of extrinsic and intrinsic forces. In the present study, we employed the selection pressure of density dependent competitive interactions according to Witting's equations (Witting, 2000) as driver for a discrete spatiotemporal model of the green oak leaf roller (Tortrix viridana). The model was successfully parameterised to rebuild the cyclic population dynamics of an empirical data set of a 30-year leaf roller monitoring in Russia. Our analysis focussed on the role of herbivore mortality and host plant food quality, which have a significant effect on T. viridana population dynamics. An additional egg or larvae mortality lowers population density and can lead to selection pressures that favour individuals with higher growth rate. This increased population growth rate can not only compensate the additional mortality, but also can lead to higher average moth abundances in subsequent generations. Furthermore, we analysed the effect of inter- and intraspecific variation in host plant quality on herbivore population dynamics and the spatial distribution of abundance and defoliation patterns. We found significant effects of the qualitative composition of a trees neighbourhood on the herbivore population of the respective tree. Also, the patchy damage patterns observable in reality have been reproduced by the present model. The applicability of the model approach and the putative genetic processes underlying Witting's model are discussed.     

Is flower scent influencing host plant selection of leaf-galling sawflies (Hymenoptera, Tenthredinidae) on willows?

Though it is known that flower scent not only attracts pollinators but also herbivores, little is known about the importance of flower scent on the distribution of leaf herbivores among individuals within a plant species. In this study we determined the distribution of galls induced by the sawfly Pontania proxima (Serville 1823) (Hymenoptera, Tenthredinidae, Nematinae) on flowering and non-flowering representatives of several clones belonging to Salix fragilis and S. × rubens (Salicaceae). Further, amounts and composition of scent of flowering and non-flowering twigs were compared (dynamic headspace-gas chromatography–mass spectrometry, DHS-GC–MS), and a scent sample collected from flowering twigs of S. fragilis was tested by coupled gas chromatography and electroantennographic detection (GC-EAD) on the antennae of P. proxima females. The results show that the presence of flower catkins on plants led to a higher degree of allocation with galls, but the number of galls differed not between flowering and non-flowering plants. The DHS-GC–MS analyses revealed that the total amount of flower scent emitted per flowering twig is about 90 times higher than the scent emitted by a non-flowering twig. Further, several compounds were emitted only by flowering but not by non-flowering twigs. In the GC-EAD analyses, antennae consistently responded not only to green leaf volatiles, but also to compounds emitted only by the flowers (e.g. 1,4-dimethoxybenzene). These flower scent compounds are suggested to affect the host plant choice by attracting more sawflies from the distance to flowering plants compared to non-flowering plants. The EAD-active compounds emitted from vegetative plant parts are assumed to act as long-distance signals especially when flowers are absent on host plants, e.g. during the oviposition period of the second generation of P. proxima.     

Current indirect fitness benefits associated with philopatry in juvenile prairie voles

Summary Prairie vole (Microtus ochrogaster) family groups were examined to determine possible indirect fitness benefits from the presence of juvenile helpers at the natal nest. The reproductive performance of family groups retaining two juveniles was compared with that of families in which all juveniles were removed when the subsequent litter was born. Litter sizes at birth of the second litter and pup survival rates were the same in the two treatments. Offspring quality was affected by the presence of juveniles however. Pups reared with juveniles weighed 13% more at weaning than pups reared without juveniles. Pups also opened their eyes sooner when juveniles were present. Differences in growth and development may have been affected by the amount of time pups were alone in the nest since pups in families with juveniles were left alone less frequently than were pups without juveniles. Maternal behavior patterns were not affected by the presence of juveniles. In contrast, fathers in families with juveniles spent more time in non-parental behaviors such as feeding, drinking, and foraging. In families with large litters, mothers delivered a subsequent litter sooner if juveniles were present. Subsequent litter sizes were the same in both treatments. Overall, both infants and parents benefited from the presence of juveniles, suggesting that helping may enhance the helper's indirect fitness in multiple ways.     

Mother really knows best: host choice of adult phytophagous insect females reflects a within-host variation in suitability as larval food

Non-random distribution patterns of specialized phytophagous insects on their hosts may depend on intraspecific differences in plant tissue quality, including nutrients and secondary compounds. Secondary compounds are involved in plant resistance, but are also important for the recognition and acceptability of plants as resources by specialized insects. If individuals within a plant species vary in their content of such secondary substances, there may also be qualitative differences between them. In such cases, natural selection will favor insects with the ability to distinguish and prefer the more suitable plants. In Sweden, the leaf beetle Gonioctena linnaeana Schrank (Coleoptera, Chrysomelidae) is highly specialized on one host, the native willow Salix triandra L (Salicaceae). Field observations reveal that some host plants in a population harbor many feeding larvae, causing severe defoliation, whereas neighboring plants may have few or no feeding larvae. Our hypothesis is that the distribution pattern of G. linnaeana larvae in this population results from qualitative differences between individual host plants in combination with the ability of G. linnaeana females to distinguish between plants that are suitable and not suitable for offspring performance. We examine whether larval survival differs depending on diet and whether the content of secondary chemical compounds explains female preference. Based on the higher survival rate of larvae reared on leaves from preferred hosts, we conclude that G. linnaeana females have evolved a behavior that maximizes offspring performance and thus positively affects female fitness. A chemical survey of the plants indicates that luteolin-7-glucoside and an unidentified flavonoid are important for separating the preferred from the non-preferred plants.