Host plant, host plant chemistry and the polyembryonic parasitoid Copidosoma sosares: indirect effects in a tritrophic interaction

Host plant identity and host plant chemistry have often been shown to influence host finding and acceptance by natural enemies but comparatively less attention has been paid to the tritrophic effects of host plant and host plant chemistry on other natural enemy fitness correlates, such as survivorship, clutch size, body size, and sex ratio. Such studies are central to understanding both the selective impact of plants on natural enemies as well as the potential for reciprocal selective impact of natural enemies on plant traits. We examined the effects of host plant and host plant chemistry in a tritrophic system consisting of three apiaceous plants (Pastinaca sativa, Heracleum sphondylium and H. mantegazzianum), the parsnip webworm (Depressaria pastinacella) and the polyembryonic parasitic wasp Copidosoma sosares. All of these plants produce furanocoumarins, known resistance factors for parsnip webworms. Furanocoumarin concentrations were correlated neither with the presence nor the number of webworms on a given plant. Concentrations of two furanocoumarins were negatively associated with C. sosares fitness correlates: isopimpinellin with the likelihood that a given webworm would be parasitized and xanthotoxin with both within‐brood survivorship (of all‐male and mixed‐sex broods) and clutch size. Brood sex ratio and body sizes of individual wasps were not correlated with furanocoumarin chemistry. Because additive genetic variation exists in P. sativa for furanocoumarin chemical traits, these are subject to selection by webworms through herbivory. Third trophic level selective impacts on furanocoumarin traits may include selection for reduced production of those chemicals that affect parasitoid survivorship yet do not influence host plant choice by the herbivore. That such might be the case is suggested by patterns of furanocoumarin production in populations of P. sativa with different histories of infestation; in the Netherlands, where parasitism rates of webworms by C. sosares are high, plants produce lower levels of all linear furanocoumarins and proportionately less isopimpinellin than do midwestern U.S. populations of P. sativa, where natural enemies of the webworm are effectively absent.     

CONSTRAINTS ON CHEMICAL COEVOLUTION: WILD PARSNIPS AND THE PARSNIP WEBWORM

The parsnip webworm (Depressaria pastinacella) and the wild parsnip (Pastinaca sativa) together represent a potentially “coevolved” system in that throughout their ranges the plant has relatively few other herbivores and the insect has virtually no other hosts. Individual wild parsnip plants within a central Illinois population vary in their content and composition of furanocoumarins, secondary compounds with insecticidal properties. Half-sib and parent-offspring regression estimates of the heritability of furanocoumarins demonstrate that this variation is genetically based. Wild parsnip plants also vary in their resistance to damage by the parsnip webworm, which feeds on flowers and developing seeds. In an experimental garden, seed production in the primary umbel ranged from 0 to 1,664 seeds among individuals, and mean seed production of half-sib families ranged from 3.7 seeds to 446.0 seeds. Approximately 75% of the variation in resistance among half-sib families to D. pastinacella was attributable to four furanocoumarin characteristics—resistance is positively related to the proportion of bergapten and the amount of sphondin in seeds, and negatively related to the amount of bergapten and the proportion of sphondin in leaves. Each of the four resistance factors had significant heritability. Thus, resistance in wild parsnip to the parsnip webworm is to a large extent chemically based and genetically controlled. Genetic correlations among fitness and resistance characters, however, tend to limit coevolutionary responses between herbivore and plant. In greenhouse plants protected from herbivory, several of the resistance factors have negative genetic correlations with potential seed production. Ostensibly, highly resistant plants in the absence of herbivory would be at a competitive disadvantage in the field. The selective impact of the herbivore is also limited in this population by a negative genetic correlation among resistance factors. Selection to increase one resistance factor (e.g., the proportion of bergapten in the seed) would at the same time decrease the amount of a second resistance factor (e.g., the amount of sphondin in the seed). The wild parsnip and the parsnip webworm, then, appear to have reached an evolutionary “stalemate” in the coevolutionary arms race.     

Phenotype matching in wild parsnip and parsnip webworms: causes and consequences

According to the geographic mosaic theory of coevolution, selection intensity in interactions varies across a landscape, forming a selection mosaic; interaction traits match at coevolutionary hotspots where selection is reciprocal and mismatch at coldspots where reciprocity is not a factor. Chemical traits play an important role in the interaction between wild parsnip (Pastinaca sativa) and the parsnip webworm (Depressaria pastinacella). Furanocoumarins, produced as plant defenses, are detoxified by the webworms by cytochrome P450 monooxygenases; significant additive genetic variation exists for both furanocoumarin production in the plant and detoxification in the insect, making these traits available for selection. To test the hypothesis that differences in selection intensity affect the distribution of coevolutionary hotspots and coldspots in this interaction, we examined 20 populations of webworms and wild parsnips in Illinois and Wisconsin that varied in size, extent of infestation, proximity to woods (and potential vertebrate predators), and proximity to a chemically distinct alternate host plant, Heracleum lanatum (cow parsnip). Twelve of 20 populations displayed phenotype matching between plant defense and insect detoxification profiles. Of the eight mismatched populations, a logistic regression model related matching probability to two predictors: the presence of the alternate host and average content of xanthotoxin (one of the five furanocoumarins produced by P. sativa). The odds of mismatching were significantly increased by the presence of the alternate host (odds ratio = 15.4) and by increased xanthotoxin content (odds ratio = 6.053). Parsnips growing near cow parsnip displayed chemical phenotypes that were chemically intermediate between cow parsnip and parsnips growing in isolation. Rapid phenotype matching in this system is likely due in part to differential mortality every season; larvae transferred to a plant 30 m or more from the plant on which they developed tended to experience increased mortality over larvae transferred to another umbel on the same plant on which they had developed, and plant populations that mismatched in 2001 displayed a change in chemical phenotype distribution from the previous year. Trait mixing through gene flow is also a likely factor in determining mismatch frequency. Populations from which webworms were eradicated the previous year were all recolonized; in three of seven of these populations, infestation rates exceeded 90%. Our findings, consistent with the geographic mosaic theory, suggest that the presence of a chemically distinct alternate host plant can affect selection intensity in such a way as to reduce the likelihood of reciprocity in the coevolutionary interaction between wild parsnip and the parsnip webworm.     

Chemical barriers to adaptation by a specialist herbivore

Summary Depressaria pastinacella, the parsnip webworm (Lepidoptera: Oecophoridae), feeds throughout eastern North America on Pastinaca sativa (wild parsnip) and few other species. The assumption that specialist herbivores such as the parsnip webworm are adapted to hostplant chemistry, and are therefore unaffected by chemical variation in hostplants, was tested. Flower buds from plants grown first in the greenhouse and then in the field were fed to ultimate instar webworms. Plant phenotype had a significant effect on virtually all webworm food utilization parameters. While nutritional factors (i.e., nitrogen content) were correlated with approximate digestibility, two constituents of the flowers — bergapten and xanthotoxin, both linear furanocoumarins — independently accounted for a significant amount of variation in food utilization indicies. The physiological effects of these furanocoumarins were confirmed in artificial diet experiments. Despite the fact that the two most important furanocoumarins in parsnip flowers relative to webworm feeding and growth are isomers, differing only in the positioning of a methoxy substituent, they have different physiological effects; while xanthotoxin in general has no effect on growth, bergapten decreases growth and digestibility of the diet. These results underscore the need in studies of plant-animal interactions to examine individual chemical components rather than classes of compounds.     

Reduced Aggressive Behavior: A Benefit of Silk-Spinning in the Parsnip Webworm, Depressaria pastinacella (Lepidoptera: Oecophoridae)

The parsnip webworm, Depressaria pastinacella, spins a silken web within the umbels of its host plant, the wild parsnip Pastinaca sativa, and aggressively defends this web against conspecifics. We first established experimentally that the number of aggressive interactions between caterpillars with their webs removed was significantly higher than for webworms with intact webs. In order to determine whether web-spinning acts to divide food resources and reduce aggressive interactions, we measured relative weight gain and total silk production of parsnip webworms isolated from one another, grouped together with webbing undisturbed, and grouped together with webbing removed daily. Parsnip webworms isolated from one another and therefore unable to engage in aggressive interactions attained the highest pupal weights and spun the smallest amount of silk; caterpillars with webs removed daily and therefore with frequent aggressive interactions until territories were reestablished had the lowest pupal weights and spun the greatest quantity of silk. Our findings indicate that, for the parsnip webworm, constructing a silken web reduces aggressive encounters among conspecifics.     

Optimal defence, kin conflict and the distribution of furanocoumarins among offspring of wild parsnip

The factors influencing the allocation of chemical defences to plant offspring have largely been unexplored, conceptually and experimentally. Because evolutionary interactions between maternal plants and their progeny can affect resource allocation patterns among sibling offspring, we suggest that kin conflict as well as herbivore–plant interaction theories need to be considered to predict chemical defence allocation patterns. Optimal defence theory predicts that maternal plants should defend more heavily those offspring in which resources have been disproportionately invested. In contrast, kin conflict theory predicts that natural selection will favour genotypes that can compete successfully for maternal defences irrespective of their quality, even at the expense of the fitness of siblings and the maternal plant. Evidence for these defence patterns were evaluated by examining the allocation of furanocoumarins to seeds of the wild parsnip (Pastinaca sativa, Apiaceae). Furanocoumarins are toxins that are localized within the oil tubes of the maternal tissues of seeds. We evaluated the role of offspring investment (endosperm mass) and seed genotype on furanocoumarin allocation by mating an array of pollen donors with pollen recipients. Furanocoumarins were found to be positively correlated with endosperm mass on one side of the seed, a result consistent with optimal defence theory; however, on the other side of the seed, furanocoumarin content was influenced by seed genotype and was unrelated to endosperm mass. These effects varied with maternal plant. Further experiments demonstrated that nearly 80% of furanocoumarin production occurs after pollination, when fertilization products are active. Although the amount of furanocoumarin influenced by the seed genotype is small relative to the total quantity in the seed, these furanocoumarins are the first line of defence against important predators, such as the parsnip webworm, Depressaria pastinacella (Lepidoptera: Oecophoridae). We found that parsnip webworm larvae were able to discriminate among genotypes within an inflorescence. In line with previous studies, these results suggest that a genotype's ability to influence furanocoumarin defence may affect its probability of survival. We conclude that the distribution of defences among plant offspring in wild parsnip is probably influenced by competition among seed genotypes that conflicts with maternal optimal defence. This revised version was published online in July 2006 with corrections to the Cover Date.     

Evolution of tolerance in an invasive weed after reassociation with its specialist herbivore

The interaction between the European wild parsnip Pastinaca sativa and its coevolved florivore the parsnip webworm Depressaria pastinacella, established in North America for over 150 years, has resulted in evolution of local chemical phenotype matching. The recent invasion of New Zealand by webworms, exposing parsnips there to florivore selection for the first time, provided an opportunity to assess rates of adaptive response in a real‐time experiment. We planted reciprocal common gardens in the USA and NZ with seeds from (1) US populations with a long history of webworm association; (2) NZ populations that had never been infested and (3) NZ populations infested for 3 years (since 2007) or 6 years (since 2004). We measured impacts of florivory on realized fitness, reproductive effort and pollination success and measured phenotypic changes in infested NZ populations relative to uninfested NZ populations to determine whether rapid adaptive evolution in response to florivory occurred. Irrespective of country of origin or location, webworms significantly reduced plant fitness. Webworms reduced pollination success in small plants but not in larger plants. Although defence chemistry remained unchanged, plants in infested populations were larger after 3–6 years of webworm florivory. As plant size is a strong predictor of realized fitness, evolution of large size as a component of florivore tolerance may occur more rapidly than evolution of enhanced chemical defence.     

Patterns of furanocoumarin production and insect herbivory in a population of wild parsnip (Pastinaca sativa L.)

Summary Seasonal changes in the distribution and abundance of furanocoumarins in wild parsnip, Pastinaca sativa (Umbelliferae), were examined in a population of plants in Tompkins County, New York. Xanthotoxin, imperatorin and bergapten (linear furanocoumarins) occur in all above-ground parts of the plant; in addition, angelicin and sphondin (angular furanocoumarins) occur in umbels of some individuals. Total furanocoumarin content, as measured by percent dry weight, is greatest in reproductive parts, particularly buds and seeds; variation in concentrations between plants is greatest in vegetative structures (e.g., leaves).Within the plant, the distribution of furanocoumarins is significantly correlated with nitrogen, as opposed to biomass, allocation. In that nitrogen is often a factor limiting the plant growth, furanocoumarins appear to be allocated in proportion to plant tissue value; reproductive structures, obvious contributors to plant fitness, contain over ten times the amount of nitrogen and furanocoumarin contained in vegetative structures such as senescent leaves.Stepwise multiple regression analysis revealed that generalized insect herbivores tend to feed on plants or plant parts low in furanocoumarin content and, correspondingly, low in nitrogen content. Parsnip specialists, notably Depressaria pastinacella (Lepidoptera: Oecophoridae), feed exclusively on umbels, plant parts rich in nitrogen and furanocoumarins; furanocoumarin number and content in fact account for over 60% of the variance in number of umbel feeders. These patterns conform with previous determinations of the toxicological properties of furanocoumarins. Nitrogen is known to affect growth rate, fecundity, longevity and survivorship of insect herbivores; by tolerating or detoxifying furanocoumarins, D. pastinacella can consume plant tissues containing significantly greater amounts of nitrogen than tissues consumed by generalist feeders. That the presence of D. pastinacella on individual plants is correlated with the number of furanocoumarins present is consistent with the hypothesis that parsnip specialists use angular furanocoumarins as host recognition cues.     

Impact of reassociation with a coevolved herbivore on oviposition deterrence in a hostplant

Although selection by herbivores for increased feeding deterrence in hostplants is well documented, selection for increased oviposition deterrence is rarely examined. We investigated chemical mediation of oviposition by the parsnip webworm (Depressaria pastinacella) on its principal hostplant Pastinaca sativa to determine whether ovipositing adults choose hostplants based on larval suitability and whether hostplants experience selection for increased oviposition deterrence. Webworms consume floral tissues and florivory selects for increased feeding deterrents; moths, however, oviposit on leaves of pre-bolting plants. Exclusive use of different plant parts for oviposition and larval feeding suggests oviposition should select for increased foliar deterrents. Recent webworm colonization of New Zealand (NZ) allowed us to assess phenotypic changes in foliar chemicals in response to webworm oviposition. In a common garden experiment, we compared NZ populations with and without a history of infestation from 2004 to 2006 for changes in leaf chemistry in response to oviposition. Three leaf volatiles, cis- and trans-ocimene, and β-farnesene, elicit strong responses in female moth antennae; these compounds were negatively associated with oviposition and are likely oviposition deterrents. Leaf β-farnesene was positively correlated with floral furanocoumarins that deter florivory; greater oviposition on plants with low floral furanocoumarins indicates that moths preferentially oviposit on parsnips most suitable for larval growth. Unlike florivory, high oviposition on leaves did not lower plant fitness, consistent with the fact that NZ parsnip foliar chemistry was unaffected by 3–6 years of webworm infestation. Thus, in this system, selection by ovipositing moths on foliar chemistry is weaker than selection by larvae on floral chemistry.     

Geographic variation in performance of a widespread generalist insect herbivore

1. Generalist herbivores are often widespread and occur in a variety of environments. Due to their broad distribution, it is likely that some populations of generalists will encounter host plants with geographic variation in traits that could affect the herbivore's growth and survival (i.e. performance). However, the geographic pattern of performance has rarely been studied for generalists, especially across large geographic ranges. 2. This study used one of the most generalist herbivore species known, the fall webworm (Hyphantria cunea Drury 1773, Erebidae, Lepidoptera), to experimentally test how the performance of a local population of fall webworms varies with increasing geographic distance of the host plant population from the local herbivore population. Specifically, a transplant experiment was used to compare the performance of one fall webworm population feeding on its local host plants with its performance on host populations from two other locations, 1300 and 2600 km away. 3. It was found that fall webworms performed better on their local host plant populations than on populations from other regions, with performance at its lowest when reared on hosts of the same species from the farthest region. It was also found that local fall webworms do not perform well on hosts commonly used by fall webworms at the other two, more distant sites. 4. This study helps to elucidate how the performance of generalist herbivores varies along their geographic range and suggests possible local adaptation to different sets of hosts across sites.     

Effects of florivory on floral volatile emissions and pollination success in the wild parsnip

In addition to reducing fitness by consuming reproductive structures, florivores may also reduce plant fitness by altering interactions with pollinators. To date, the effects of florivore activity on the volatile profile of flowers and subsequent attractiveness to pollinators have not been extensively investigated. In this study, we had three specific objectives: to determine the impact of florivory by the parsnip webworm Depressaria pastinacella on the floral volatile profile of the wild parsnip Pastinaca sativa, to ascertain the mechanisms by which florivory changes the volatile profile, and to estimate the consequences of florivory on visitation by pollinators and eventual seed set. An overall indirect effect of webworms on seed set, that is, the effect of infestation on pollination success, was not detected. However, this overall lack of indirect effect masks the heterogeneity among individual plants. For seven of 14 plants examined, pollination success was altered by webworms, and in four of these plants the alteration in pollination success was consistent with webworm-altered visitation. Webworms significantly altered floral fragrance, in particular causing disproportionate increases in the emissions of octyl esters. Additionally, volatiles from webworm frass, which contains large amounts of the octyl ester metabolite n-octanol, may alter the floral fragrance in ways that change attractiveness of flowers to pollinators. This study suggests that the effects of florivores on plant fitness are not limited to the removal of floral units but may also involve alterations in floral volatile composition, through damage-induced release and detoxification of particular constituents, that affect visitation and pollination success. Handling Editor: Steve Johnson. An erratum to this article can be found at     

Heritability estimates for octyl acetate and octyl butyrate in the mature fruit of the wild parsnip

The aliphatic esters octyl acetate and octyl butyrate occur as major components of essential oils in the vittae, or oil tubes, of the wild parsnip (Pastinaca sativa). We determined phenotypic variation and narrow-sense heritabilities of these octyl esters in wild parsnip fruits from 30 maternal families. The mean octyl acetate content was 1.56 microg/mg dry fruit (0.08-5.51 microg/mg dry fruit) and the mean octyl butyrate content was 4.28 microg/mg dry fruit (1.28-14.22 microg/ mg dry fruit). Narrow-sense heritabilities for each ester's content were calculated by analysis of half-sib families (HS) and parent-offspring regression (OP). Heritabilities were 0.389 (HS) and 0.654 (OP) for octyl acetate and 0.670 (HS) and 0.626 (OP) for octyl butyrate. The amounts of the esters were phenotypically correlated with each other and with the linear furanocoumarins bergapten and xanthotoxin, phototoxic compounds that co-occur in the vittae with the esters. Ester amounts were not genetically correlated, indicating that these compounds could respond independently to selection pressures. These octyl esters may serve as carrier solvents that enhance penetration of these furanocoumarins into herbivore integuments and gut walls.     

Physiological price of an induced chemical defense: photosynthesis, respiration, biosynthesis, and growth

A recurring theme in defense allocation theories is that defenses are costly. Most studies that attempt to quantify a cost of defense seek to establish a trade-off between a component of plant fitness and the level of a constitutive defense. Such estimates are ambiguous because they cannot discount the cost of traits that are correlated with defense but are not themselves defensive. We examined the effects of damage-induced synthesis of furanocoumarins, known defense compounds, on the growth of wild parsnip. Plants that had 2% of their leaf area removed accumulated 8.6% less total biomass and 14% less root biomass than intact plants over a 4-week period. We also found that this small amount of leaf damage significantly reduced net photosynthetic rates 0.5 h after damage; the effect was temporary, as photosynthetic rates were no longer significantly different after 48 h. Lastly, we found that increases in respiration rates associated with damage coincided spatially and temporally with increases in furanocoumarin production, and that respiration increases were phenotypically correlated with furanocoumarin production. When damage-induced changes in furanocoumarin content and respiration rates were expressed in glucose equivalents and compared, the energetic cost of furanocoumarin production (12.6 μg glucose cm⁻²) accounted for all of the increase in respiration (12.0 μg glucose cm⁻²). A comparison of other secondary compounds in damaged and intact leaflets revealed that myristicin, a furanocoumarin synergist, is the only other compound aside from furanocoumarins that is inducible. The inducible defense system of wild parsnip thus appears to involve a small subset of secondary compounds. Synthesis of these compounds is tightly linked to damage-induced rates of respiration. Because the negative impact that damage had on the rate of net photosynthesis was short-lived, the impact of damage on growth observed in this study was likely due to the cost of furanocoumarin synthesis elicited by damage rather than the loss of photosynthetic tissue caused by damage. Received: 4 April 1996 / Accepted: 29 August 1996     

Parthenocarpic fruits in wild parsnip: Decoy defence against a specialist herbivore

Summary Parthenocarpy, the production of fruits without viable seeds, is a widespread phenomenon in plants. While failure to effect pollination or fertilization is often cited as the cause of parthenocarpy, this explanation alone is inadequate to explain why plants produce, maintain and further develop fruits. Wild parsnips (Pastinaca sativa) frequently produce parthenocarpic fruit. When parsnip webworms (Depressaria pastinacella), specialist feeders on wild parsnip, were given choices between normal fruit and parthenocarpic fruit, they exhibited a strong preference for parthenocarpic fruit. However, on parthenocarpic fruit, insects fed less efficiently and grew more slowly than insects fed normal fruit. Parthenocarpic fruits, then, may act as decoys that divert herbivores away from fruits that contain plant offspring.     

Furanocoumarins in seeds of wild and cultivated parsnip

The furanocoumarin content of ripe seeds of seven cultivars of Pastinaca sativa was compared to that of ripe seeds from a naturalized population in central Illinois. Five furanocoumarins were identified and quantified in the intact seeds. While seeds of wild and cultivated plants contain furanocoumarin components in identical proportions, seeds from wild plants contain almost three times the quantity of furanocoumarins on a dry weight basis as do seeds of cultivars.     

Generalist and specialist host–parasitoid associations respond differently to wild parsnip (Pastinaca sativa) defensive chemistry

1. Plant defensive chemistry is predicted to have a more negative effect on generalist herbivores and their parasitoids than on specialist herbivores and their parasitoids. 2. This prediction was examined by comparing the effects of the wild parsnip (Pastinaca sativa L.) toxin, xanthotoxin, on a generalist herbivore–parasitoid association [the cabbage looper, Trichoplusia ni Hübner, and its polyembryonic parasitoid, Copidosoma floridanum (Ashmead)] and a specialist herbivore–parasitoid association [the parsnip webworm, Depressaria pastinacella (Duponchel), and its polyembryonic parasitoid, Copidosoma sosares (Walker)]. 3. Copidosoma floridanum brood sizes were smaller and experienced lower survivorship when reared in a host feeding on an artificial diet containing a low concentration of xanthotoxin. No T. ni hosts, parasitised or unparasitised, survived on a diet high in xanthotoxin. In contrast, C. sosares brood size and survivorship were unaffected by the presence of low levels of xanthotoxin in the host diet. Copidosoma sosares experienced reduced brood size and survivorship only when its host consumed a diet containing 15 times the level of xanthotoxin as the diet adversely affecting its congener. 4. The differences in response to xanthotoxin exhibited by C. floridanum and C. sosares are explained partly by a differential reduction in host quality and partly by differential exposure to xanthotoxin in host haemolymph. Unlike D. pastinacella, T. ni experienced reduced pupal weight and survivorship and prolonged developmental time on a low‐xanthotoxin diet. More xanthotoxin passed unmetabolised into the haemolymph of T. ni than into the haemolymph of D. pastinacella.     

Allelic variation in the Depressaria pastinacella CYP6AB3 protein enhances metabolism of plant allelochemicals by altering a proximal surface residue and potential interactions with cytochrome P450 reductase

CYP6AB3v1, a cytochrome P450 monooxygenase in Depressaria pastinacella (parsnip webworm), is highly specialized for metabolizing imperatorin, a toxic furanocoumarin in the apiaceous host plants of this insect. Cloning and heterologous expression of CYP6AB3v2, an allelic variant identified in D. pastinacella, reveals that it metabolizes imperatorin at a rate (V(max) of 10.02 pmol/min/pmol of cytochrome P450 monooxygenase (P450)) significantly higher than CYP6AB3v1 (V(max) of 2.41 pmol/min/pmol) when supplemented with even low levels of cytochrome P450 reductase. Comparisons of the NADPH consumption rates for these variants indicate that CYP6AB3v2 utilizes this electron source at a faster rate than does CYP6AB3v1. Molecular modeling of the five amino acid differences between these variants and their potential interactions with P450 reductase suggests that replacement of Val(92) on the proximal face of CYP6AB3v1 with Ala(92) in CYP6AB3v2 affects interactions with P450 reductase so as to enhance its catalytic activity. Allelic variation at this locus potentially allows D. pastinacella to adapt to both intraspecific and interspecific variation in imperatorin concentrations in its host plants.     

HOST-PATHOGEN INTERACTIONS IN NATURAL POPULATIONS OF LINUM MARGINALE AND MELAMPSORA LINI: II. LOCAL AND REGIONAL VARIATION IN PATTERNS OF RESISTANCE AND RACIAL STRUCTURE

Spatial variation in the resistance structure of Linum marginale (wild flax) populations to the rust fungus Melampsora lini, and in the racial structure of this pathogen, was investigated by sampling 10 populations distributed throughout the Kosciusko National Park, New South Wales, Australia. Considerable differences were found among populations in the structure of both host and pathogen. Host populations were divided into three broad categories: (1) populations susceptible to all testing races; (2) populations containing a strictly limited number of resistant phenotypes; and (3) populations with a considerable diversity of resistant phenotypes. The pathogen populations also showed a range of diversity. The major differences between these populations were determined by the occurrence and frequency of four common races of pathogen (races A, E, K, and N). These differences were apparent both at a regional spatial scale (over the 100 km separation of the most distant populations) and at a local scale where major differences were detected between two populations only 300 m apart. The distribution of the four common races of M. lini was consistent with the hypothesis that a fitness cost was associated with unnecessary virulence. In general, however, differences in the structure of pathogen populations from genetically very similar host populations implied that, in addition to host resistance genes, other evolutionary forces are also important in determining the genetic structure of individual pathogen populations.     

Abundance trumps quality: bi‐trophic performance and parasitism risk fail to explain host use in the fall webworm

Factors including host abundance, quality, and the degree to which hosts provide enemy‐free space (EFS) may drive host plant choice by phytophagous insects. Herbivores may also experience fitness tradeoffs among hosts, promoting polyphagy. The fall webworm Hyphantria cunea is a dietary generalist that feeds on a broad array of trees across its geographic range. Here, we investigate the drivers of host tree use by the fall webworm in Connecticut (CT) and Maryland (MD). Neither caterpillar performance nor EFS was associated with the frequency with which host trees were used, and no tradeoff between host quality and EFS was identified. Vegetation surveys adjacent to host trees showed that at both localities, host use was non‐random with respect to tree species, and that the main predictor of use among suitable host trees was host tree abundance. This suggests that webworms are under selection to reduce search time for oviposition sites. Although we did not detect a tradeoff between host plant quality and availability in MD, we did identify that tradeoff in CT. This disparity amid otherwise similar patterns of host use between CT and MD may be explained by the relative rarity of high quality hosts in CT compared to MD. Our results illustrate that geographic mosaics in patterns of host use may arise in the absence of local adaptation if host use is based upon availability rather than host plant attributes.     

Host Genetics and Environment Drive Divergent Responses of Two Resource Sharing Gall-Formers on Norway Spruce: A Common Garden Analysis

A central issue in the field of community genetics is the expectation that trait variation among genotypes play a defining role in structuring associated species and in forming community phenotypes. Quantifying the existence of such community phenotypes in two common garden environments also has important consequences for our understanding of gene-by-environment interactions at the community level. The existence of community phenotypes has not been evaluated in the crowns of boreal forest trees. In this study we address the influence of tree genetics on needle chemistry and genetic x environment interactions on two gall-inducing adelgid aphids (Adelges spp. and Sacchiphantes spp.) that share the same elongating bud/shoot niche. We examine the hypothesis that the canopies of different genotypes of Norway spruce (Picea abies L.) support different community phenotypes. Three patterns emerged. First, the two gallers show clear differences in their response to host genetics and environment. Whereas genetics significantly affected the abundance of Adelges spp. galls, Sacchiphantes spp. was predominately affected by the environment suggesting that the genetic influence is stronger in Adelges spp. Second, the among family variation in genetically controlled resistance was large, i.e. fullsib families differed as much as 10 fold in susceptibility towards Adelges spp. (0.57 to 6.2 galls/branch). Also, the distribution of chemical profiles was continuous, showing both overlap as well as examples of significant differences among fullsib families. Third, despite the predicted effects of host chemistry on galls, principal component analyses using 31 different phenolic substances showed only limited association with galls and a similarity test showed that trees with similar phenolic chemical characteristics, did not host more similar communities of gallers. Nonetheless, the large genetic variation in trait expression and clear differences in how community members respond to host genetics supports our hypothesis that the canopies of Norway spruce differ in their community phenotypes.