Host plant preference based on salicylate chemistry in a willow leaf beetle (Chrysomela aeneicollis)

Summary Chrysomela aeneicollis (Coleoptera: Chrysomelidae) uses salicin from its host plant (Salix spp.) to produce a defensive secretion, salicylaldehyde. Because it requires salicin for this secretion, I predicted that C. aeneicollis should be attracted to willows which possess salicin and other salicylates. To test this prediction, I determined the host-plant preferences of C. aeneicollis among four potential hosts which occur in the Sierra Nevada range of eastern California. These species have very different salicylate chemistries but do not differ in nutritional quality for C. aeneicollis. In oviposition-preference tests, gravid females showed no preference between a salicylate-poor species, S. lutea, and a salicylate-rich species, S. orestera. However in feeding-choice tests, both larvae and adults preferred S. orestera over S. lutea. This preference was not affected by the species on which the larvae were reared. In other feeding tests, adults preferred S. orestera over two medium-salicylate species, S. boothi and S. geyeriana, regardless of which host species they had been feeding on in nature. In a final feeding test, adults were stimulated to feed by salicin itself. In nature, the relative abundances of C. aeneicollis adults and egg clutches among these species correspond to the adult feeding preference in the laboratory. Additionally, multiple regression analyses showed that adult abundance was not related to among-clone differences in leaf toughness or nutritional quality, but rather to salicin content and plant size. Thus for C. aeneicollis, both laboratory and field results demonstrate a preference for salicylate-rich willows which is partly responsible for the increased level of attack on them.     

Enemy-free space? Host preference and larval performance of a willow leaf beetle

To examine whether enemy-free space is an important factor determining the host utilization pattern of a leaf beetle Plagiodera versicolora, we investigated the relationship between adult preference and offspring performance on three co-occurring willow species, Salix sachalinensis, S. miyabeana and S. integra. Salix sachalinensis was by far the most preferred host plant of feeding adults, while both S. miyabeana and S. integra were rarely fed upon. The fact that most oviposition was observed on S. sachalinensis also suggested that P. versicolora preferred S. sachalinensis to other willows for oviposition. This adult preference did not correspond well to patterns of larval performance on the three willow species in the absence of enemies. Higher survivorship, shorter developmental time and larger adult size were achieved on S. sachalinensis and S. miyabeana than on S. integra. Performance as indicated by female adult size and development time on S. miyabeana were higher than on S. sachalinensis. In the presence of enemies, however, the survivorship of first-instar larvae on S. miyabeana was much lower than on other willows. Adults of P. versicolora apparently avoided S. miyabeana as an oviposition and feeding host and preferred S. sachalinensis as an enemy-free space. This was not because larvae had poorer performance on S. miyabeana, but because predation pressure on eggs and early instar larvae was more severe on S. miyabeana.     

Predator avoidance as a function of flocking in the sexually dichromatic Hawaii akepa

Hypotheses for joining a mixed-species bird flock consider each species as a single unit. In sexually dichromatic birds, differential conspicuousness between the sexes may result in differences in vigilance for predators. Aspects of the predator avoidance and foraging enhancement hypotheses for the selective value of joining a mixed-species flock were assessed for the strongly sexually dichromatic Hawaii akepa (Loxops coccineus coccineus). There was support for the primary predictions of the predator avoidance hypothesis: vigilance levels decreased with increasing group size, and with membership in a flock, but only for brightly colored adult males. There was little support for the hypothesis that the primary benefit of joining a mixed-species flock is to enhance foraging efficiency through local enhancement.     

Factors contributing to the seasonal occurrence of the willow leaf beetle Plagiodera versicolora

We conducted a 3-year study of a natural population of the willow leaf beetle Plagiodera versicolora (Coleoptera: Chrysomelidae) on a river bank of the Inukami River, Shiga, central Japan, where four willow species (Salix chaenomeloides, S. eriocarpa, S. integra, and S. serissaefolia) occur sympatrically. Our survey showed that: (1) at the study site, the abundance of P. versicolora greatly varied among years and among willow species; (2) adult abundance changed seasonally with species-specific patterns on different willow species; and (3) the dispersal-settlement of adults had the most pronounced effects on the seasonal population growth rate of P. versicolora. Factors affecting these results were discussed.     

Predator induced life-history shifts in a freshwater cladoceran

Summary Life-history theory predicts that maturity and resource allocation patterns are highly sensitive to selective predation. Under reduced adult survival, selection will favour genotypes capable of reproducing earlier, at a smaller size and with a higher reproductive effort. When exposed to water that previously held fish, (size selective predators which prefer larger Daphnia), individuals of Daphnia hyalina reproduced earlier, at a smaller size and had a higher reproductive investment. Hence the prey was able to switch its life history pattern in order to become less susceptible to predation by a specific predator. The cue that evokes the prey response is a chemical released by the predator.     

Host-plant effects on larval survival of a salicin-using leaf beetle Chrysomela aeneicollis Schaeffer (Coleoptera: Chrysomelidae)

Several species of willow leaf beetles use hostplant salicin to produce a defensive secretion that consists of salicylaldehyde. Generalist arthropod predators such as ants, ladybird beetles, and spiders are repelled by this secretion. The beetle larvae produce very little secretion when they feed on willows that lack salicylates, and salicin-using beetles prefer salicylate-rich willows over salicylate-poor ones. This preference may exist because the larvae are better defended against natural enemies on salicylate-rich willows. If this is true, the larvae should survive longer on those willows in nature. However, this prediction has not been tested. I determined the larval growth and survival of Chrysomela aeneicollis (Coleoptera: Chrysomelidae) on five willow species (Salix boothi, S. drummondiana, S. geyeriana, S. lutea, and S. orestera). These species differed in their salicylate chemistries and in leaf toughness but not in water content. The water content varied among the individual plants. Larval growth of C. aeneicollis did not differ among the five species in the laboratory, but it varied among the individual plants and it was related to the water content. In the field, C. aeneicollis larvae developed equally rapidly on the salicylate-poor S. lutea and on the salicylate-rich S. orestera. Larval survival was greater on S. orestera than on S. lutea in one year (1986), but there was no difference between them during three succeeding years. In another survivorship experiment, larval survival was low on the medium-salicylate S. geyeriana, but high on the salicylate-poor S. boothi and on S. orestera. Larval survival in the field was related to the larval growth and water content that had been previously measured in the laboratory. These results showed that the predicted relationship between the host plant chemistry and larval survival did not usually exist for C. aeneicollis. One possible reason for this was that the most important natural enemies were specialist predators that were unaffected by the host-derived defensive secretion. One specialist predator, Symmorphus cristatus (Hymenoptera: Eumenidae), probably caused much of the mortality observed in this study. I discuss the importance of other specialist predators to salicin-using leaf beetles.     

Predator avoidance behaviour in the buffy-headed marmoset,Callithrix flaviceps

The predator avoidance behaviour of a free-ranging group of buffy-headed marmosets,Callithrix flaviceps, was recorded in detail during the course of a long-term study of behavioural ecology at the Fazenda Montes Claros, southeastern Brazil. Four distinct patterns of predator avoidance behaviour, each with specific vocalisations, were recognised and are described here. The selection and use of sleeping sites by the study group are also described. An analysis of the records indicates that these small monkeys are generally most vulnerable to predation by aerial raptors. Variations in the frequency of alarm calls also indicate that the marmosets tend to be more vigilant at higher levels in the forest and when the leaf cover is less extensive. The implications of group size and social structure for both the evolution and the efficacy of the anti-predator behaviour of marmosets are also discussed.     

Patch-exploitation patterns in an egg parasite of elm leaf beetle

Eggs of elm beetle,Pyrrhalta luteola (Muller), normally occur in masses which can be viewed as discrete host patches in space and time. Analysis of>800 egg masses from 3 field sites in northern California revealed that the number of eggs/mass varied from 2 to>40 and that the imported egg parasiteTetrastichus gallerucae (Fonsc.) exploited a relatively large proported of the egg masses at certain times. The spatial relationship between parasitization and number of eggs/egg mass was assessed in 2 ways-i.e., for exploited masses only and for all masses combined (exploited+nonexploited). Percent parasitization was density independent in most cases for both data sets. It is suggested that both methods of analysis can provide patterns which are relevant to biological control of insect pests.      

Slow larval growth on a suboptimal willow results in high predation mortality in the leaf beetle Galerucella lineola

The slow growth/high mortality hypothesis predicts that herbivorous insects feeding on suboptimal host plants are subjected to higher predation mortality owing to the longer time spent in the vulnerable juvenile stages compared with conspecifics feeding on optimal plants. We tested this hypothesis for the willow-feeding leaf beetle Galerucella lineola raised on one suitable (Salix viminalis) and one unsuitable (S. dasyclados) willow species as well as on plants from an interspecific cross between the two species. Cohorts of larvae raised on caged plants (protected from enemies) and uncaged plants (exposed to enemies) were monitored daily throughout larval development in two consecutive years. Larvae raised on S. viminalis developed faster, grew larger and survived better than those raised on S. dasyclados. The suitability of the hybrid plants was intermediate to that of their parents. Our results strongly support the slow growth/high mortality hypothesis. In both years, total predation during the larval period was higher on S. dasyclados than on S. viminalis. Furthermore, the daily predation rate (i.e. the proportion of larvae preyed upon per day) was higher on S. dasyclados than on S. viminalis. When hybrid plants were included in the analysis total predation was positively correlated with both larval development time and daily predation rate. We suggest that high predation on beetles on low-quality plants is the combined result of their longer development time and elevated daily predation rate. The results are discussed in relation to the evolution of host plant selection and the paradox of sublethal plant defenses.     

The effects of chrysomelid beetle grazing and plant competition on the growth of Rumex obtusifolius

Summary The interaction between grazing by Gastrophysa viridula and interspecific plant competition was investigated for Rumex obtusifolius growing in the field. During an eight-month growing season non-competing R. obtusifolius grew larger than competing plants although herbivore loads (beetle productivity per unit leaf area) were similar for competing and non-competing plants. However, grazing significantly reduced the growth of R. obtusifolius only in competing plants thus demonstrating a synergism between plant competition and invertebrate herbivory. The relevance of these findings for the distribution of plant species is discussed.     

Plant stress and insect performance: cottonwood,ozone and a leaf beetle

Summary Leaf area consumption rates, development rates, survivorship, and fecundity of the imported willow leaf beetle (Plagiodera versicolora Laich) were examined on two clones of eastern cottonwood which were previously exposed to ozone or charcoal-filtered air. P. versicolora consumed more ozone treated foliage, but were more fecund when reared on charcoal-filtered air treated plants. Beetle development rates and survivorship were not significantly different on treated and control cottonwoods. We concluded that: 1) Ozone fumigation of cottonwood reduced foliage quality, and the reproductive success and overall performance of P. versicolora. 2) increased foliage consumption by beetles was probably a mechanism compensating for decreases in foliage quality. 3) Reductions in beetle fecundity were due to an initial reduction in oviposition rates. 4) Beetle feeding preference did not correlate with the suitability of foliage for beetle performance. These results are discussed in relation to the impact of air pollution on plant-insect interactions.     

Dynamics of leaf minerals,bdleaf area,and biomass from hardwoods intensively grown on a peat bog

Summary Water willow and grey alder were grown on a raised sphagnum bog in central Sweden. The stands were intensively treated by daily irrigation and fertilization during the growing period in order to improve site fertility. After a 2-year establishment period high production rates were achieved in willow stands, 0.8 kg stem dry weight m^–2 year^–1 on current plus one (C+1) year old shoots. In these stands the canopy was closed with a leaf area index (LAI) that peaked at approximately 7. The canopy in the alder stand did not close during the initial 3 years of growth and the measured production rate was relatively low, at approximately 0.4 kg dry weight m^–2 year ^–1 in the last year. The leaf nitrogen content was 3%–4% of dry weight during the summer and the other studied mineral elements were in almost optimal proportion to nitrogen. This was considered to be an effect of the intensive fertilization regime. Above-ground production close to maximum yield was attained in the prevailing conditions of soil, climate and biomass partitioning.     

Impact of epidermal leaf mining by the aspen leaf miner (Phyllocnistis populiella) on the growth, physiology, and leaf longevity of quaking aspen

The aspen leaf miner, Phyllocnistis populiella, feeds on the contents of epidermal cells on both top (adaxial) and bottom (abaxial) surfaces of quaking aspen leaves, leaving the photosynthetic tissue of the mesophyll intact. This type of feeding is taxonomically restricted to a small subset of leaf mining insects but can cause widespread plant damage during outbreaks. We studied the effect of epidermal mining on aspen growth and physiology during an outbreak of P. populiella in the boreal forest of interior Alaska. Experimental reduction of leaf miner density across two sites and 3 years significantly increased annual aspen growth rates relative to naturally mined controls. Leaf mining damage was negatively related to leaf longevity. Leaves with heavy mining damage abscised 4 weeks earlier, on average, than leaves with minimal mining damage. Mining damage to the top and bottom surfaces of leaves had different effects on physiology. Mining on the top surface of the leaf had no significant effect on photosynthesis or conductance and was unrelated to leaf stable C isotope ratio (delta(13)C). Mining damage to the bottom leaf surface, where stomata are located, had significant negative effects on net photosynthesis and water vapor conductance. Percent bottom mining was positively related to leaf delta(13)C. Taken together, the data suggest that the primary mechanism for the reduction of photosynthesis by epidermal leaf mining by P. populiella is the failure of stomata to open normally on bottom-mined leaves.     

Components of resistance of leaf rust (Melamspora laricii epitea Kleb./Melampsora ribesii-viminalis Kleb.) in Salix viminalis L.

Selection on partial resistance components, namely latent period, development rate of uredia and number and size of uredia, has been suggested as a means to achieve durable resistance. Three experiments were carried out in growth chambers to assess the impact of environmental and genetic factors on these components in the Salix-Melampsora host-pathogen system. They confirmed the environmental ability of the Melampsora-Salicaceae relationship and provided no definite answer on the possibility of attaining durable resistance through selection on partial resistance components. On the one hand, there is a large amount of heritable variation among clones for most components; on the other hand, all components were extremely sensitive to environmental conditions and isolates, making the development of standard selection methods difficult.     

Sex differentiation in postnatal growth rate: a test in a wild boar population

Summary In wild boar individual growth rate is linear between 0.5 and 6 months after birth, based on successive body weight measurements. Contrary to expectation for a dimorphic and polygynous mammal like wild boar, no sexual dimorphism in growth rate could be detected between 0.5 and 6 months. We argue that high total maternal invesment in offspring due to large litter size and/or strong selection for early reproduction in this population with a short generation time could explain this absence of early differentiation in postnatal growth rate according to offspring sex.     

Chlorogenic acid as an antiherbivore defence of willows against leaf beetles

In the leaves of 13 Finnish willow species, the content of a phenolic, chlorogenic acid, was found to vary from 0 up to 18 mg g^–1 D.W. Effects of pure chlorogenic acid on insect feeding behaviour were tested using four common leaf beetle species which are in the field mainly found on willows with low-chlorogenic acid leaves. One species, Lochmaea capreae L., was invariably deterred by pure chlorogenic acid applied in naturally occurring concentrations on the willow leaves. Accordingly, in 2-choice laboratory feeding trials L. capreae was found to prefer low-chlorogenic acid leaves of four willow species over high-chlorogenic acid leaves of Salix pentandra L. and S. myrsinifolia Salisb. When presented on the leaves of S. phylicifolia L, pure chlorogenic acid inhibited also the feeding by Phratora polaris Sp.-Schn. Instead, chlorogenic acid had no significant effect on Ph. polaris when it was presented on the leaves of another willow S. cinerea L. In laboratory, Ph. polaris did not show general preference for willow species with low chlorogenic acid content in their leaves. Thus, the response of Ph. polaris to chlorogenic acid seems to depend on the plant species. Apparently variation in other traits such as leaf hairyness may easily override the potential effect of chlorogenic acid content on Ph. polaris. To two other leaf beetle species, Galerucella lineola F. and Plagiodera versicolora Laich., chlorogenic acid is an ineffective deterrent even at unnaturally high concentrations. In laboratory, G. lineola and P. versicolora did not prefer willows with low chlorogenic acid content in their leaves. Thus, among four studied leaf beetle species, only L. capreae seems to be clearly affected by this phenolic. Therefore, overall importance of chlorogenic acid as a defence against willow-feeding leaf beetles appears to be very limited.     

The quorum-sensing system in a plant bacterium Mesorhizobium huakuii affects growth rate and symbiotic nodulation

Mesorhizobium huakuii is a free-living bacterium which is capable of establishing a specific symbiotic relationship with Astragalus sinicus, an important winter green manure widely used in Eastern Asia, allowing for nitrogen fixing during this process. Previous studies demonstrate that M.␣huakuii produces quorum-sensing molecules at high cell density and quorum sensing plays a role in biofilm formations. In this study, we isolated and characterized two quorum-sensing deficient mutants in M. huakuii. Analysis of the flanking region of transposon insertions indicated that autoinducer synthase related genes are not homologous to acyl homoserine lactone (AHL) synthase genes that are shared among many Gram-negative bacteria, but related to peptide synthesis, indicating that M. huakuii quorum-sensing signals are distinct from AHLs. Compared with the wild-type strains, these quorum-sensing deficient mutants promoted their growth rate and were defective in nodule formation on host plants, indicative of a critical role of quorum sensing in M.␣huakuii during the host–bacterium symbiotic interaction.Yijing Gao and Zengtao Zhong contributed equally to this work.     

Early spring leaf out enhances growth and survival of saplings in a temperate deciduous forest

Saplings of many canopy tree species in winter deciduous forests receive the major portion of their light budget for their growing season prior to canopy closure in the spring. This period of high light may be critical for achieving a positive carbon (C) gain, thus contributing strongly to their growth and survival. This study of saplings of Aesculus glabra and Acer saccharum in Trelease Woods, Illinois, USA, tested this hypothesis experimentally by placing tents of shade cloth over saplings during their spring period of high light prior to canopy closure in three consecutive years. Leaf senescence began 16 days (year 0) and 60 days (year 1) earlier for shaded A. glabra saplings than control saplings. No change in senescence occurred for A. saccharum. The annual absolute growth in stem diameter of both species was negligible or negative for shaded saplings, but positive for control saplings. Only 7% of the shaded A. glabra saplings were alive after 2 years, while all control saplings survived for 3 years; only 20% of the shaded A. saccharum saplings survived for 3 years, while 73% of control saplings were alive after the same period. Early spring leaf out is a critical mechanism that allows the long-term persistence of saplings of these species in this winter deciduous forest. Studies and models of C gain, growth, and survival of saplings in deciduous forests may need to take into account their spring phenology because saplings of many species are actually “sun” individuals in the spring prior to their longer period in the summer shade.     

Interference competition between ladybird beetle adults (Coleoptera: Coccinellidae): effects on growth and reproductive capacity

Cage experiments were used to determine the minimum number of Aphis fabae Scopoli that females of the Azorean native ladybird beetle Coccinella undecimpunctata L. require per day to achieve maximum fecundity and to assess the effects of interference competition from conspecific and heterospecific [Harmonia axyridis (Pallas)] adult ladybirds on its growth and reproduction. The number of A. fabae at which females of C. undecimpunctata ate most aphids, achieved maximum reproduction and maintained their body weight, was 200. The presence of H. axyridis adults, unlike the presence of C. undecimpunctata, significantly affected the reproductive numerical response of C. undecimpunctata even though the resource was not in short supply. The results suggest that interference competition with heterospecifics may adversely affect the reproductive capacity of C. undecimpunctata. Based on previous results and those reported here, we suggest that the introduction of H. axyridis could endanger the population abundance and/or result in the competitive displacement of C. undecimpunctata from the Azorean ecosystem.     

Host race formation in the meadowsweet and strawberry feeding leaf beetle Galerucella tenella

We investigated host race formation in Galerucella tenella, a leaf beetle which feeds preferentially on meadowsweet (Rosaceae) in its natural habitats, but has become an important pest on strawberry (Rosaceae) in agricultural areas. Beetles from two isolated sites (Skeppsvik in Sweden and Solf in Finland) were compared with respect to preference and performance. At Skeppsvik the beetles were found feeding on meadowsweet, while at Solf large populations only develop in strawberry plantations, despite the presence of meadowsweet.In reciprocal field transplantations and laboratory bioassays, beetles from both sites discriminated against their foreign host, using their natal plant to a significantly higher degree for both egg laying and feeding, but with one interesting exception, namely that beetles from Solf increased their use of and even preferred meadowsweet for feeding in the laboratory. This increased use of meadowsweet by adult Solf beetles in the laboratory (without conspecific competitors) may be attributed to a density-dependent strategy, whereby mothers avoid pre-emptying the nutritional resources of the plants that will host their offspring in cases where no competitors are present.Larval fitness did not differ significantly between host plants, although larval survival of Solf beetles was halved when reared on their non-natal host plant (meadowsweet). Although beetles from both sites preferred to oviposit on their local host plant, our results provide little evidence that this presumed adaptation has to do with the nutritional quality of the plants involved. Instead, we suggest that other factors associated with the plants, such as enemy-free space and resistance to drought may be more important selective agents, shaping host preference in the field.