Nutrition as a facilitator of host‐race formation: the shift of a stem‐boring beetle to a gall host

1. The importance of host‐race formation to herbivorous insect diversity depends on the likelihood that successful populations can be established on a new plant host. A previously unexplored ecological aid to success on a novel host is better nutritional quality. The role of nutrition was examined in the shift of the stem‐boring beetle Mordellistena convicta to fly‐induced galls on goldenrod and the establishment there of a genetically distinct gall host race. 2. First, larvae of the host race inhabiting stems of Solidago gigantea were transplanted into stems and galls of greenhouse‐grown S. gigantea plants. At the end of larval development, the mean mass of larvae transplanted to galls was significantly greater than the mass of larvae transplanted to stems, indicating a likely nutritional benefit during the shift. This advantage was slightly but significantly diminished when the gall‐inducing fly feeding at the centre of the gall died early in the season. Additionally, there was a suggestion of a trade‐off in the increased mortality of smaller beetle larvae transplanted into galls. 3. In a companion experiment, S. gigantea gall‐race beetle larvae were likewise transplanted to S. gigantea stems and galls. Besides the expected greater mass in galls, the larvae also exhibited adaptations to the gall nutritional environment: larger inherent size, altered tunnelling behaviour, and no diminution of mass pursuant to gall‐inducer mortality. 4. In a third line of inquiry, chemical analyses of field‐collected S. gigantea plants revealed higher levels of mineral elements important to insect nutrition in galls as compared with stems.     

Gall Flies, Inquilines, and Goldenrods: A Model for Host-race Formation and Sympatric Speciation

Host shifts and subsequent host-race formation likely play amore common role in the speciation of herbivorous insects thanhas generally been recognized. Our studies of the interactionsof goldenrod host plants (Solidago: Compositae), the gall flyEurosta solidaginis (Diptera: Tephritidae), and the stem- andgall-boring Mordellistena convicta (Coleoptera: Mordellidae)provide behavioral, ecological, and genetic evidence of insecthost races that may represent incipient species formed via sympatricspeciation. Eurosta solidaginis has developed genetically differentiatedand reproductively isolated host races that are associated withthe ancestral host Solidago altissima and the derived host S.gigantea. Conventional wisdom suggests that shifts even to closelyrelated host plants are limited by host preferences or the inabilityto utilize a chemically and developmentally distinct host. However,our preliminary work with Eurosta from S. gigantea implies thathost choice and gall induction do not deter a shift to S. canadensis.The galling of Solidago by Eurosta created a new resource thathas led to a subsequent host range expansion by the stem-boringbeetle. Mordellistena convicta from stems and galls are geneticallydistinct and likely shifted from stems to galls. Beetles fromS. altissima versus S. gigantea galls exhibit assortative matingand higher preference for and/or performance on their natalhost. The present-day distributions of the Eurosta host racesand their behavioral isolating mechanisms do not suggest thatgeographic isolation was required for their formation; ratherthese characteristics suggest a sympatric mode of differentiation.Our findings lend credence to recent assertions that sympatricspeciation may be an important source of biodiversity.     

Phenotypic plasticity and similarity among gall morphotypes on a superhost,Baccharis reticularia (Asteraceae)

Understanding factors that modulate plant development is still a challenging task in plant biology. Although research has highlighted the role of abiotic and biotic factors in determining final plant structure, we know little of how these factors combine to produce specific developmental patterns. Here, we studied patterns of cell and tissue organisation in galled and non‐galled organs of Baccharis reticularia, a Neotropical shrub that hosts over ten species of galling insects. We employed qualitative and quantitative approaches to understand patterns of growth and differentiation in its four most abundant gall morphotypes. We compared two leaf galls induced by sap‐sucking Hemiptera and stem galls induced by a Lepidopteran and a Dipteran, Cecidomyiidae. The hypotheses tested were: (i) the more complex the galls, the more distinct they are from their non‐galled host; (ii) galls induced on less plastic host organs, e.g. stems, develop under more morphogenetic constraints and, therefore, should be more similar among themselves than galls induced on more plastic organs. We also evaluated the plant sex preference of gall‐inducing insects for oviposition. Simple galls were qualitative and quantitatively more similar to non‐galled organs than complex galls, thereby supporting the first hypothesis. Unexpectedly, stem galls had more similarities between them than to their host organ, hence only partially supporting the second hypothesis. Similarity among stem galls may be caused by the restrictive pattern of host stems. The opposite trend was observed for host leaves, which generate either similar or distinct gall morphotypes due to their higher phenotypic plasticity. The Relative Distance of Plasticity Index for non‐galled stems and stem galls ranged from 0.02 to 0.42. Our results strongly suggest that both tissue plasticity and gall inducer identity interact to determine plant developmental patterns, and therefore, final gall structure.     

The influence of host plant variation and intraspecific competition on oviposition preference and offspring performance in the host races of Eurosta solidaginis

1. A series of experiments was conducted to measure the impact of plant genotype, plant growth rate, and intraspecific competition on the oviposition preference and offspring performance of the host races of Eurosta solidaginis (Diptera: Tephritidae), a fly that forms galls on Solidago altissima and Solidago gigantea (Asteraceae). Previous research has shown that both host races prefer to oviposit on their own host plant where survival is much higher than on the alternate host plant. In this study, neither host race showed any relationship between oviposition preference and offspring performance in choosing among plants of their natal host species. 2. The larval survival of both host races differed among plant genotypes when each host race oviposited on its natal host species. In one experiment, altissima host race females showed a preference among plant genotypes that was not correlated with offspring performance on those genotypes. In all other experiments, neither the altissima nor gigantea host race demonstrated a preference for specific host plant genotypes. 3. Eurosta solidaginis had a preference for ovipositing on rapidly growing ramets in all experiments, however larval survival was not correlated with ramet growth rate at the time of oviposition. 4. Eurosta solidaginis suffered high mortality from intraspecific competition in the early larval stage. There was little evidence, however, that females avoided ovipositing on ramets that had been attacked previously. This led to an aggregated distribution of eggs among ramets and strong intraspecific competition. 5. There was no interaction among plant genotype, plant growth rate, or intraspecific competition in determining oviposition preference or offspring performance.     

ONE HOST SHIFT LEADS TO ANOTHER? EVIDENCE OF HOST‐RACE FORMATION IN A PREDACEOUS GALL‐BORING BEETLE

Abstract.— We show that a predator, the tumbling flower beetle Mordellistena convicta (Coleoptera: Mordellidae), has formed host races in response to a host-plant shift and subsequent host-race formation by its prey, the gall-inducing fly Eurosta solidaginis (Diptera: Tephritidae). This fly has formed two host races, one that induces stem galls on the ancestral host plant, Solidago altissima (Compositae), and another that induces stem galls on the closely related S. gigantea . We found that subpopulations of M. convicta that attack E. solidaginis galls on the different host plants have significantly different emergence times and, although slight, these allochronic differences are consistent across a range of temperatures. More importantly, we found that beetles assortatively mate according to their natal host plants, and female M. convicta preferentially attack and/or their offspring have higher survival in galls on natal host plants. Our data suggest that subpopulations of M. convicta that attack E. solidaginis galls on S. altissima and S. gigantea have formed host races. This is one of the first studies to demonstrate that a host shift and subsequent host-race formation by an herbivorous insect may have resulted in subsequent diversification by one of its natural enemies.     

Host‐race formation: promoted by phenology,constrained by heritability

Host‐race formation is promoted by genetic trade‐offs in the ability of herbivores to use alternate hosts, including trade‐offs due to differential timing of host‐plant availability. We examined the role of phenology in limiting host‐plant use in the goldenrod gall fly (Eurosta solidaginis) by determining: (1) whether phenology limits alternate host use, leading to a trade‐off that could cause divergent selection on Eurosta emergence time and (2) whether Eurosta has the genetic capacity to respond to such selection in the face of existing environmental variation. Experiments demonstrated that oviposition and gall induction on the alternate host, Solidago canadensis, were the highest on young plants, whereas the highest levels of gall induction on the normal host, Solidago gigantea, occurred on intermediate‐age plants. These findings indicate a phenological trade‐off for host‐plant use that sets up the possibility of divergent selection on emergence time. Heritability, estimated by parent–offspring regression, indicated that host‐race formation is impeded by the amount of genetic variation, relative to environmental, for emergence time.     

Divergent host plant preference causes assortative mating between sympatric host races of the ladybird beetle,Henosepilachna diekei

Divergent host preference (i.e. host fidelity) plays a significant role in the speciation process in phytophagous insects. However, how and to what extent this divergence reduces gene flow between populations has rarely been measured. Here, we estimated the intensity of assortative mating caused solely by host fidelity in two host races of the phytophagous ladybird beetle Henosepilachna diekei, specialized on Mikania micrantha (Asteraceae) and Leucas lavandulifolia (Lamiaceae) in West Java, Indonesia. These host races mated randomly in the absence of host plants under laboratory conditions, but demonstrated nearly complete assortative mating in field cages with the two host plants, by spending almost all of their time on their respective host plants. The frequency of assortative mating in the field cages was not affected drastically by host plant patch structure. These results suggest that fidelity to the different host plants yields directly almost complete reproductive isolation between the host races by limiting the habitat on the respective host plant. In addition, the high host fidelity also ensures female oviposition on the original host plant. As larvae cannot survive on non‐host plants, a positive association between female oviposition preference and larval performance on the host plant on which the beetles are specialized will further facilitate the evolution of host fidelity. © 2013 The Linnean Society of London, Biological Journal of the Linnean Society, 2013, 110 , 606–614.     

Behavioural,ecological and genetic evidence confirm the occurrence of host‐associated differentiation in goldenrod gall‐midges

Host‐associated differentiation (HAD) is considered a step towards ecological speciation and an important mechanism promoting diversification in phytophagous insects. Although the number of documented cases of HAD is increasing, these still represent only a small fraction of species and feeding guilds among phytophagous insects, and most reports are based on a single type of evidence. Here we employ a comprehensive approach to present behavioural, morphological, ecological and genetic evidence for the occurrence of HAD in the gall midge Dasineura folliculi (Diptera: Cecidomyiidae) on two sympatric species of goldenrods (Solidago rugosa and S. gigantea). Controlled experiments revealed assortative mating and strong oviposition fidelity for the natal‐host species. Analysis of mitochondrial DNA showed an amount of genetic divergence between the two host‐associated populations compatible with cryptic species rather than host races. Lower levels of within‐host genetic divergence, gall development and natural‐enemy attack in the S. gigantea population suggest this is the derived host.     

The effects of host race, gender, and host plant distribution on alighting behavior, mating, and oviposition in Eurosta solidaginis

Eurosta solidaginis Fitch (Diptera: Tephritidae) has formed host races on Solidago altissima L. and Solidago gigantea Ait. (Asteraceae), and reproductive isolation between these host races is brought about in part by host‐associated assortative mating. Any non‐assortative mating creates the potential for gene flow between the populations, and we investigated the conditions that favored non‐assortative mating. We hypothesized that the frequency of non‐assortative mating would be influenced by differences in the behaviors of the host races and sexes and by the presence and pattern of distribution of the two host species. To test these hypotheses, we caged flies on four combinations of 32 potted host plants: all S. altissima, all S. gigantea, and cages with both host species arranged in either two pure species blocks or randomly dispersed. We recorded the number of flies of each host race that alighted on each host species and the frequency of mating within and between the host races. Males of both host races were observed on plants more frequently than females. Flies of the host race from S. gigantea (gig flies) were observed on plants in greater absolute numbers, and they mated more frequently than flies of the host race from S. altissima (alt flies). In all treatments, gig flies of both sexes were found on non‐natal host plants significantly more frequently than alt flies, and gig females showed a weaker preference for their host species than did gig males or alt flies of either gender for their respective natal hosts. Assortative mating predominated in all treatments, and flies from each host race mated more frequently in cages containing their own host plant. The frequency of non‐assortative mating varied among treatments, with the matings between alt ♀ × gig ♂ being more common in the pure S. altissima treatment and the gig ♀ × alt ♂ being more frequent in the pure S. gigantea and random treatments. Matings between gig ♂ × alt ♀ were more common overall than the reciprocal mating, because gig males were more active in pursuing matings and in alighting on the non‐natal host plant than alt flies. Non‐assortative matings were more frequent in the random than in the block treatments, but this difference was not significant. Because of strong selection against oviposition into the alternate host, we hypothesized that host plant distribution would not affect oviposition preference. We tested this hypothesis by examining the oviposition behavior of naïve, mated females in two treatments in which both host species were present: either arranged in blocks or randomly dispersed. Females oviposited only into their natal host, regardless of host plant distribution.     

Divergence of Eurosta solidaginis in response to host plant variation and natural enemies

We tested the hypothesis that forest and prairie populations of the gall-inducing fly, Eurosta solidaginis, have diverged in response to variation in selection by its host plant Solidago altissima, and its natural enemies. A reciprocal cross infection design experiment demonstrated that fly populations from the prairie and forest biomes had higher survival on local biome plants compared to foreign biome host plants. Flies from each biome also had an oviposition preference for their local plants. Each fly population induced galls of the size and shape found in their local biome on host plants from both biomes indicating a genetic basis to the differences in gall morphology. Solidago altissima from the prairie and forest biomes retained significant morphological differences in the common garden indicating that they are genetically differentiated, possibly at the subspecies level. The populations are partially reproductively isolated as a result of a combination of prezygotic isolation due to host-associated assortative mating, and postzygotic isolation due to low hybrid survival. We conclude that E. solidaginis is undergoing diversifying selection to adapt to differences between prairie and forest habitats.     

Endophytic insect communities of two prairie perennials (Asteraceae: Silphium spp.)

Little is known of the biology of most insects that are endemic to prairie ecosystems of North America, with the exception of large and conspicuous species. In particular, species that are sequestered within plant tissues are commonly overlooked. In this paper, we assess the biodiversity of endophytic insects that inhabit stems of Silphium laciniatum L. and S. terebinthinaceum Jacquin (Asteraceae), endemic plants of tallgrass prairies. Endophytic herbivores, gall wasps Antistrophus rufus Gillette and A. minor Gillette (Hymenoptera: Cynipidae) and stem-boring larvae of the beetle Mordellistena aethiops Smith (Coleoptera: Mordellidae) were attacked by 10 species of natural enemies. We report new host plant associations for herbivores, and new host insect associations for parasitoids. The two plant species differed significantly in their densities of gall wasps and the vertical dispersion of galls within stems. Interactions within and between trophic levels attest to the biodiversity of endophytic insect communities, and the specialized nature of these insects suggests they are highly vulnerable to habitat conservation practices that involve destruction of dead vegetation.     

Tritrophic interactions and trade-offs in herbivore fecundity on hybridising host plants

Abstract. 1. Interspecific plant hybridisation can have important evolutionary consequences for hybridising plants and for the organisms that they interact with on multiple trophic levels. In this study the effects of plant hybridisation on the abundance of herbivores and on the levels of herbivore parasitism were investigated. 2. Borrichia frutescens, B. arborescens, and their hybrid (B. × cubana) were censused for Asphondylia borrichiae galls and Pissonotus quadripustulatus plant hoppers in the Florida Keys. Levels of egg parasitism were determined by dissecting parental and hybrid stems and galls for herbivore and parasite eggs and larvae. Stem toughness and gall size are plant‐mediated modes of protection from parasitism and these were also measured. For gall midges, fly size was measured as an estimate of fecundity. 3. Field censuses indicated that herbivore abundances varied on hybrid hosts relative to parent plant species and that the different herbivore species exhibited different patterns of abundance. Asphondylia borrichiae gall numbers followed the additive pattern of abundance while P. quadripustulatus numbers most closely resembled the dominance pattern. 4. Parasitism of P. quadripustulatus eggs was high on B. frutescens and the hybrids, and low on B. arborescens, which also had significantly tougher stems. Asphondylia borrichiae suffered the highest levels of parasitism on B. frutescens, the host plant which produced the smallest galls. On B. arborescens, which produced the largest galls, levels of A. borrichiae parasitism were lowest. Both parasitism and gall size were intermediate on the hybrid plants. Galls on B. arborescens and hybrid plants produced significantly smaller flies then those from B. frutescens suggesting that, when selecting hosts from among parent species and hybrids, gall flies may face a trade‐off between escape from natural enemies and maximising fecundity.     

Testing the nutrition hypothesis for the adaptive nature of insect galls: does a non‐adapted herbivore perform better in galls?

Abstract.  1. The nutrition hypothesis for the adaptive nature of galls states that gall-inducing insects control the nutrient levels in galls to their own benefit. Although the nutrition hypothesis is widely accepted, there have been few empirical tests of this idea.
2. A novel method is presented for testing the nutrition hypothesis that links manipulation of gall nutrient levels by the gall inducer to herbivore performance. The effects of adaptation and nutritional advantage are separated by using a herbivore that is adapted to a host plant susceptible to galling but one which never enters the gall environment.
3.  Hellinsia glenni (Cashatt), a plume moth (Pterophoridae) and one of its host plants provide an excellent system for testing the nutrition hypothesis because H. glenni larvae feed internally on the relatively nutrient-poor stems of a goldenrod, Solidago gigantea , but do not venture into the nutrient-rich galls induced on that plant by a tephritid fly, Eurosta solidaginis . The nutrition hypothesis was tested by transplanting early-instar H. glenni larvae into galls and stems of S. gigantea to determine if the larvae transplanted to galls would perform better compared with those larvae transplanted to stems.
4. The results support the nutrition hypothesis for the adaptive nature of galls. Hellinsia glenni achieved greater final mass in the gall environment compared with the final mass larvae achieved in the stem environment. There was also evidence that the quality of gall tissue is controlled by the gall inducer, which has not been previously demonstrated for mature E. solidaginis galls.     

Plant phenology-mediated indirect effects: The gall midge opens the phenological window wider for a leaf beetle

We examined whether larvae of the gall midge Rabdophaga rigidae (Diptera: Cecidomyiidae) can modify the seasonal dynamics of the density of a leaf beetle, Plagiodera versicolora (Coleoptera: Chrysomelidae), by modifying the leaf flushing phenology of its host willow species, Salix serissaefolia and Salix eriocarpa (Salicaceae). To test this, we conducted field observations and a laboratory experiment. The field observations demonstrated that the leaf flushing phenology of the willows and the seasonal dynamics of the beetle density differed between shoots with stem galls and shoots without them. On galled shoots of both willow species, secondary shoot growth and secondary leaf production were promoted; consequently, leaf production showed a bimodal pattern and leaf production periods were 1 to 2 months longer than on non‐galled shoots. The adult beetle density on galled shoots was thus enhanced late in the season, and was found to change seasonally, synchronizing with the production of new leaves on the host willow species. From the results of our laboratory experiment, we attributed this synchrony between adult beetle density and willow leaf flush to beetles’ preference to eat new leaves rather than old. Indeed, beetles consumed five times more of the young leaves when they were fed both young and old leaves. These results indicate that stem galls indirectly enhance the adult beetle density by enhancing food quality and quantity late in the beetle‐feeding season. We therefore conclude that midge galls widen the phenological window for leaf beetles by extending the willows’ leaf flush periods.     

Oviposition site preference and developmental performance of a gall‐inducing leafhopper on galled and non‐galled host plants

Oviposition preferences of herbivorous insects affect offspring performance. Both positive and negative links between oviposition preference and offspring performance have been reported for many species. A gall‐inducing leafhopper, Cicadulina bipunctata Melichar (Hemiptera: Cicadellidae), feeds on various Poaceae plants and induces galls of enhanced nutritional value for their offspring. Although gall induction by C. bipunctata improves nymphal performance, the oviposition preference of females between galled and non‐galled host plants is still unclear. In this paper, the nymphal performance and oviposition and feeding‐site preference of C. bipunctata were investigated using galled wheat, Triticum aestivum L., and non‐galled barley, Hordeum vulgare L., as host plants. The survival rate of C. bipunctata on wheat was significantly higher than on barley. In the choice test, significantly more eggs were laid into barley, whereas the number of eggs deposited on both hosts was not significantly different in the no‐choice test. The number of settling individuals per leaf area was not significantly different between wheat and barley, suggesting no clear preference for oviposition between these plants. Experience as a nymph with a growing host did not affect oviposition preference as adult female. The inconsistent correspondence between offspring performance and oviposition preference of C. bipunctata may reflect the high mobility of nymphs and/or differences in leaf area between host plants. The results indicate that the previous finding that oviposition preference and offspring performance are not always positively correlated in herbivorous insects is applicable to gall‐inducing insects.     

Do parasitoids diversify in response to host‐plant shifts by herbivorous insects?

1. For herbivorous insects, the incorporation of a novel host into the diet, and subsequent formation of distinct host associations (races), is thought to be a significant early step in the speciation process. While many studies have addressed this issue, virtually nothing is known about the evolutionary response of natural enemies to herbivore host‐race formation. 2. The hypothesis that the parasitoid wasp Eurytoma gigantea (Hymenoptera: Eurytomidae) has formed host races in direct response to the host shift and subsequent host‐race formation by its host, the gallmaker Eurosta solidaginis (Diptera: Tephritidae) was tested. Emergence time, mating preference, and female oviposition preference were determined for parasitoids derived from galls of each Eurosta host race. 3. Male and female E. gigantea overlap broadly in their emergence times from each Eurosta host race, suggesting that there is no phenological barrier to gene flow. 4. In choice experiments, female parasitoids did not mate assortatively: females that emerged from one Eurosta host race were equally likely to mate with males from either Eurosta host race. 5. Oviposition behaviour experiments revealed that female parasitoids do not prefer to oviposit on their host race of origin and that there is no overall preference for one host race, even though fitness is higher when parasitoids are reared from Eurosta galls of the Solidago gigantea host race than when reared from Eurosta galls of the Solidago altissima host race. 6. These results suggest that E. gigantea has not diverged in parallel with its host in response to the herbivore host‐plant shift. Further studies are needed before the ubiquity of this diversification mechanism can be evaluated fully.     

Abundance of Gall Midges on Poulsenia armata (Moraceae): Importance of Host Plant Size and Light Environment in Tropical Rain Forests1

We studied the effects of contrasting light environments on the relationship between the host plant size of Poulsenia armata and the abundance of two gall midges in a tropical rain forest in Veracruz, Mexico. The number and density of two gall morphs (i.e., laminar and vein‐petiole galls) were positively correlated with plant size only in trees found in the forest but not in gaps. The availability of foliar area of P. armata trees was greater in forest gaps than in the forest. The foliar area was positively correlated with the abundance of laminar galls in trees in forest sites, but not with vein‐petiole galls. We concluded that the abundance of two morphs of gall midges on P. armata was associated with host plant size only in the forest trees. Larger plants had more galls than small plants, although this relationship was affected by local light environments.     

The protein content of tissues in cynipid galls (Hymenoptera: Cynipidae): Similarities between cynipid galls and seeds

Cynipid galls are examples of induced plant development, where the gall inducer is in control of cell differentiation and morphogenesis of a new plant organ. This study concentrates on the tissues of the larval chamber common to all cynipid galls. The protein content of the inner gall tissue was compared to that of non‐gall plant tissues. We investigated three oak and two rose galls and their respective host plants. Total protein signatures of inner gall tissues were different from those of non‐gall plant tissues, and among the five galls. N‐terminal sequences were obtained for two abundant proteins from the inner gall tissues of D. spinosa and A. quercuscalicis, which were common to all galls, at 62 and 43 kDa. Database queries suggest the 62 kDa protein to be homologous to a protein disulphide isomerase (PDI), and the 43 kDa protein to be homologous to NAD‐dependent formate dehydrogenase (FDH). A naturally biotinylated protein was detected at 33 kDa during Western analyses with streptavidin. Western analyses revealed the presence of the biotinylated protein and PDI in the inner gall tissues of all five galls, while FDH was only detected in A. quercuscalicis and A. fecundator. PDI was also common to all non‐gall tissues, while FDH was not detected in non‐gall tissues, and the biotinylated protein was only detected in seeds. The proteins identified in the inner gall tissue suggest that (a) inner gall tissues in some galls are under respiratory stress, and (b) cynipid gall formation might involve the ectopic expression of seed‐specific proteins.     

Sequential radiation of unrelated organisms: the gall fly Eurosta solidaginis and the tumbling flower beetle Mordellistena convicta

Host shifts and the formation of insect-host races are likely common processes in the speciation of herbivorous insects. The interactions of goldenrods Solidago (Compositae), the gall fly Eurosta solidaginis (Diptera: Tephritidae) and the beetle Mordellistena convicta (Coleoptera: Mordellidae) provide behavioural, ecological and genetic evidence of host races that may represent incipient species forming via sympatric speciation. We summarize evidence for Eurosta host races and show that M. convicta has radiated from goldenrod stems to Eurosta galls to form host-part races and, having exploited the galler's host shift, has begun to differentiate into host races within galls. Thus, host-race formation has occurred in two interacting, but unrelated organisms representing two trophic levels, resulting in 'sequential radiation' (escalation of biodiversity up the trophic system). Distributions of host races and their behavioural isolating mechanisms suggest sympatric differentiation. Such differentiation suggests host-race formation and subsequent speciation may be an important source of biodiversity.     

Preference–performance relationship in the gall midge Rabdophaga rosaria: insights from a common‐garden experiment with nine willow clones

1. Oviposition preferences of herbivorous insects are predicted to match offspring performance on different host taxa or on conspecific host genotypes. In gall‐inducing insects, host‐plant properties such as growth rate and gall size, which are determined by plant genotype and growing conditions, may have a significant impact on offspring performance and, hence, should influence oviposition site selection. 2. The present study investigated host preference of the European rosette willow gall midge Rabdophaga rosaria (Loew) in relation to offspring success on seven clones of Salix myrsinifolia Salisb. and two naturally hybridised S. myrsinifolia × phylicifolia L. clones growing in a replicated design in an experimental field under two fertilisation regimes. For each clone, the average growth rate, number of shoot tips, and leaf and gall size were determined, and their effects on midge preference and larval survival were examined. 3. Main shoot height, number of shoot tips, and gall size were significantly affected by clone. The midges clearly preferred certain clones over the others, but preferences were not related to willow growth traits or to gall size. Survival probability was higher in large than in small galls, but females did not prefer large‐leaved clones that produced the biggest rosette galls. Midge oviposition was also uncorrelated with prior rates of leaf‐rust infection and with feeding preferences of voles and folivorous insects. 4. The weak preference–performance relationship of R. rosaria within S. myrsinifolia is probably explained by evolutionary constraints that prevent generalist insects from achieving an ability to discriminate among conspecific hosts of variable quality.