Aggregation facilitates larval growth in the neotropical nymphalid butterfly Chlosyne janais

1. Larvae of Chlosyne janais (Lepidoptera: Nymphalidae) feed gregariously as early instars on the shrub Odontonema callistachyum (Acanthaceae). During the fourth instar, aggregations break up and larvae feed as solitary individuals.
2. The hypothesis that aggregation increases growth rate was tested by raising larvae on intact plants in the field in different group sizes and measuring their daily growth.
3. There was a striking effect of group size on larval growth whereby larvae more than doubled their weight gain by feeding in large rather than small aggregations on intact plants in the field.
4. This group-feeding advantage was lost altogether if larvae were raised on excised leaves in the laboratory, suggesting that large aggregations may facilitate growth either by inducing a nutrient sink or by overwhelming an induced allelochemical response in the plant.
5. Although larval survival was higher in cages that excluded enemies than in exposed aggregations, there was no influence of group size (experimentally manipulated) on short-term survival in the field. However, there was a weak positive relationship between short-term survival and the size of naturally occurring larval aggregations in the field. These data provide mixed support for the notion that gregarious feeding promotes defence against natural enemies.
6. Although the group defence hypothesis warrants further investigation, feeding facilitation is clearly an important factor contributing to the aggregation behaviour of C. janais larvae.     

The consequences of larval aggregation in the butterfly Chlosyne lacinia

1. Females of Chlosyne lacinia (Geyer) (Lepidoptera: Nymphalidae, Melitaenae), the bordered patch butterfly, clump eggs in a few large clusters on their host plant, Helianthus annuus. Resulting larvae form sibling aggregations to at least the third instar.
2. The effect of group size on survival and development of C. lacinia larvae was tested experimentally in the field. Larvae developed faster and survived better in larger groups.
3. The effects of various predator guilds (ground-dwelling arthropods, aerial arthropods and avian predators) on survival of larvae was then tested while controlling group size. Ground-dwelling arthropods, mainly fire ants Solenopsis xyloni , reduced larval survival greatly but other solitary invertebrate and avian predators did not alter survival. Group defences and aposematism of C. lacinia larvae are probably ineffective against predatory ants that attack en masse and recruit other colony members.
4. In laboratory experiments, two possible mechanisms underlying faster development of larvae in larger groups were tested: (i) overcoming the physical toughness of host plant leaves, and (ii) social stimulus to feed. Results support the physical toughness hypothesis but not the social stimulus hypothesis.
5. Feeding in large groups by C. lacinia larvae confers multiple advantages, including protection from solitary predators and increased feeding efficiency because grouped, early-instar larvae can initiate feeding wounds on tough sunflower leaves. These advantages of larval gregariousness, coupled with reduced desiccation at the egg stage, apparently outweigh disadvantages of aggregation, such as interference and exploitative competition among larvae.     

Life-history responses of larviparous Boettcherisca formosensis (Diptera: Sarcophagidae) to larval competition for food, including comparisons with oviparous Hemipyrellia ligurriens (Calliphoridae)

Abstract. 1 The effect of larval rearing density on life-history parameters of Boettcherisca formosensis Kirner & Lopes (Sarcophagidae) was investigated. Increases in rearing density resulted in lowered larval survivorship, shortened larval development time and production of smaller, shorter-lived adults with reduced fecundity.
2. B. formosensis is larviparous. Average brood size was 17.5±1.0 (mean±M) larvae, which was much less than the average number of mature larvae inside gravid females. Females apparently produced a series of small broods, distributing their offspring over a number of carcasses.
3. Compared with the oviparous species Hemipyrellia ligurriens (Wiedemann) (Calliphoridae), B. formosensis adults were larger and longer-lived, with a longer larval development time but shorter larval feeding period. However, females had a shorter pre-reproductive period, were less fecund, and had a lower life time reproductive investment.
4. B. formosensis had lower relative performance (measured by the composite index of performance, r') than H. ligurriens over the larval rearing density range, and was more sensitive to increases in density. Although the r' values suggest that the sarcophagid may be a competitively inferior species, other features which are not included in the index (such as larvipary, short larval feeding period and spreading of offspring from a single brood among carcasses) may be of significant adaptive value to B. formosensis.     

Interactions among white spruce tannins, Bacillus thuringiensis subsp. kurstaki, and spruce budworm (Lepidoptera: Tortricidae), on larval survival, growth, and development

The interactions among white spruce, Picea glauca (Moench) Voss, purified acetone tannin extracts (hydrolyzable and condensed tannin), Bacillus thuringiensis subsp. kurstaki Cry1A(c) delta-endotoxin strain HD-73 (Btk), and spruce budworm, Choristoneura fumiferana (Clemens) (Lepidoptera: Tortricidae) on larval survival, growth, and development were investigated over the whole larval feeding period by using artificial diet supplemented with three concentrations of Btk toxins per milliliter of diet (0, 0.021, and 1.72 microg/ml) and three concentrations of foliar tannin extract (0, 8, and 15% dry mass basis). At high Btk concentration, tannin antagonized Btk potency against spruce budworm by lowering Btk-related larval mortality from 83 to 43%. At moderate Btk concentration tannin did not affect Btk potency. Host tree tannins antagonized not only the lethal effects of Btk toxin but also sublethal Btk-related impacts in terms of larval development, pupal weight, relative consumption rate, and growth rate. When alone in the diet, tannin negatively affected larval survival, growth, and development. Maximum potency of tannins against spruce budworm larvae (60% mortality) was reached at dietary concentrations corresponding to what is found in the plant (8% dry mass). The addition of Btk toxin in food containing tannin reduced percentage of larval mortality by one-third, indicating that Btk toxin can antagonize tannin potency against the insect. Development of Btk transgenic spruce trees should consider the antagonistic effect the toxin may have on the resistance conferred by tannins that have evolved naturally in spruce trees.     

Differential defoliation of Eucalyptus grandis arises from indiscriminant oviposition and differential larval survival

1 The influence of six open-pollinated families (OPFs) of Eucalyptus grandis on both the growth and development of larvae and the oviposition preference of a paropsine chrysomelid ( Paropsis atomaria ) was investigated. The OPFs had previously been identified as differing in their susceptibility to defoliation by P. atomaria in forestry progeny trials.
2 Oviposition preference for resistant and susceptible foliage was tested using binary choice tests. These tests did not demonstrate any significant preference for either resistant or susceptible open-pollinated material indicating that adult host preference for susceptible trees was not a likely cause of differential defoliation.
3 Quantification and analysis of growth and development parameters for all larval stages of P. atomaria showed that feeding on genetic material identified as resistant resulted in a significant reduction of relative growth rate of first instar larvae and an alteration to normal feeding behaviour. There was also a trend towards increased larval mortality on resistant E. grandis .
4 We argue that although the magnitude of these effects was minor, interactions with additional biotic and abiotic sources of mortality in the field have the potential, when magnified over successive generations, to result in significant variation in defoliation of host genotypes in the field.     

Temporal and spatial variations in microclimate influence the larval foraging behaviors and performance of a conifer-feeding sawfly

Herbivorous insects are often exposed to broad temporal and spatial variations in microclimate conditions within their host plants and have adapted a variety of behaviors, such as avoidance or basking, to either offset or benefit from such variation. Field experiments were carried out to investigate the influence of daily and intratree variations in microclimate on the behaviors (feeding, resting, dispersal, and hiding) and associated performance of late-instar larvae of the yellowheaded spruce sawfly, Pikonema alaskensis (Rohwer) (Hymenoptera: Tenthredinidae) within crowns of 1.25-1.5 m tall black spruce (Picea mariana [Miller] Britton Sterns Poggenburg); late instars feed on developing shoots of young spruce and are often exposed to microclimatic extremes with unknown effects on performance. Larvae fed diurnally from just after dawn (0800 h) until dusk (2000 h) and rested throughout the night, with brief periods of dispersal occurring in the morning and evening. Neither larval behavior nor abiotic conditions differed significantly between the upper and lower crowns of trees. Temperature, humidity, and solar insolation all explained >90% of variation in feeding; however, sunrise and sunset were the most likely cues influencing diurnal behavior. Most larvae (94%) fed on the bottom, shaded side of shoots, and field experiments indicated that this behavior is adaptive with respect to microclimate, probably reducing hygrothermal stress. Thus, behavioral adaptations by P. alaskensis to daily and within-shoot microclimatic variation may reduce the risk of hygrothermal stress during dispersal or feeding, while still allowing larvae to feed on the preferred and highly nutritious upper crown foliage of young spruce.     

Environmental variation and behaviour: resource availability during ontogeny and feeding performance in salamander larvae (Ambystoma texanum)

1. The effects of resource availability during ontogeny on subsequent feeding performance were investigated in larvae of the small-mouthed salamander ( Ambystoma texanum ).
2. Salamander larvae were reared individually in either high or low prey density treatments for 7 weeks prior to intermediate prey density foraging trials. Larvae from the low prey density treatment were on average 35% smaller in body size than individuals from the high prey density treatment.
3. Resource availability during development influenced larval feeding rates and altered the relationship between body size and three feeding performance measures (attack rates, capture success and feeding rates). Feeding rates in predation trials were also positively correlated with growth rate early in the larval period (until the end of week 5).
4. These results suggest that the environment to which developing organisms are exposed can have significant effects on subsequent behaviour, and that small-mouthed salamander larvae may show state-dependent changes in feeding behaviour in response to differences in long-term feeding history. Additionally, differences in feeding performance may influence the probability of survival to the adult stage for organisms that utilize ephemeral habitats.     

Environmental variation and behaviour: resource availability during ontogeny and feeding performance in salamander larvae (Ambystoma texanum)

1. The effects of resource availability during ontogeny on subsequent feeding performance were investigated in larvae of the small-mouthed salamander ( Ambystoma texanum ).
2. Salamander larvae were reared individually in either high or low prey density treatments for 7 weeks prior to intermediate prey density foraging trials. Larvae from the low prey density treatment were on average 35% smaller in body size than individuals from the high prey density treatment.
3. Resource availability during development influenced larval feeding rates and altered the relationship between body size and three feeding performance measures (attack rates, capture success and feeding rates). Feeding rates in predation trials were also positively correlated with growth rate early in the larval period (until the end of week 5).
4. These results suggest that the environment to which developing organisms are exposed can have significant effects on subsequent behaviour, and that small-mouthed salamander larvae may show state-dependent changes in feeding behaviour in response to differences in long-term feeding history. Additionally, differences in feeding performance may influence the probability of survival to the adult stage for organisms that utilize ephemeral habitats.     

Oviposition site preference and larval mortality in a leaf-mining moth

Abstract. 1. The univoltine leaf-mining moth, Lithocolletis quercus Ams., is endemic to Israel, where it spends its 10.5 month larval period feeding only in the leaves of Quercus calliprinos Webb.
2. We compared patterns of egg deposition and sources of larval mortality to test whether oviposition patterns and site preferences confer an enhanced likelihood of larval survival.
3. Dominant sources of larval mortality were premature leaf abscission and death from unknown causes, whereas predation, parasitism and intraspecific interference accounted for relatively little larval mortality.
4. Eggs, and thus mines, were aggregated among leaves of host trees even though premature leaf abscission was positively correlated with density of mines per leaf. Interference competition among larvae was the only other density-dependent mortality factor.
5. Oviposition patterns within leaves mitigated the probability of death from larval interference, and probably also from early leaf abscission.
6. Despite these density-dependent mortality factors, overall probability of larval survival to pupation was independent of initial density of mines on a leaf.
7. The long larval period allows synchrony between oviposition flights and times of predictable resource availability.     

Allee effect in larval resource exploitation in Drosophila: an interaction among density of adults, larvae, and micro-organisms

Abstract 1. Aggregation pheromones can evolve when individuals benefit from clustering. Such a situation can arise with an Allee effect, i.e. a positive relationship between individual fitness and density of conspecifics. Aggregation pheromone in Drosophila induces aggregated oviposition. The aim of the work reported here was to identify an Allee effect in the larval resource exploitation by Drosophila melanogaster, which could explain the evolution of aggregation pheromone in this species. 2. It is hypothesised that an Allee effect in D. melanogaster larvae arises from an increased efficiency of a group of larvae to temper fungal growth on their feeding substrate. To test this hypothesis, standard apple substrates were infested with specified numbers of larvae, and their survival and development were monitored. A potential beneficial effect of the presence of adult flies was also investigated by incubating a varying number of adults on the substrate before introducing the larvae. Adults inoculate substrates with yeast, on which the larvae feed. 3. Fungal growth was related negatively to larval survival and the size of the emerging flies. Although the fungal growth on the substrate was largely reduced at increased larval densities, the measurements of fitness components indicated no Allee effect between larval densities and larval fitness, but rather indicated larval competition. 4. In contrast, increased adult densities on the substrates prior to larval development yielded higher survival of the larvae, larger emerging flies, and also reduced fungal growth on the substrates. Hence, adults enhanced the quality of the larval substrate and significant benefits of aggregated oviposition in fruit flies were shown. Experiments with synthetic pheromone indicated that the aggregation pheromone itself did not contribute directly to the quality of the larval resource. 5. The interaction among adults, micro‐organisms, and larval growth is discussed in relation to the consequences for total fitness.     

Seasonal variation in the reproductive strategy of the parasitic wasp Eulophus larvarum (Hymenoptera: Chalcidoidea: Eulophidae)

ABSTRACT. 1. Eulophus larvarum (L.) (Hymenoptera: Eulophidae) is a gregarious parasitoid of lepidopterous larvae feeding on broad-leaved trees. Normally there are two generations a year.
2. Sex ratio in the spring generation of larvae is female biased. The bias is probably due to local mate competition as progeny from one brood emerge from their pupae in close proximity to each other.
3. Sex ratio in the summer generation of larvae is near equality. Local mate competition is absent as individuals from the same brood become separated during the winter.
4. Variation in sex-ratio in the spring generation is consistent with a binomial model. Variation in sex ratio in the summer generation is much greater than expected from a binomial model and there is a large proportion of single-sex broods.
5. Two hypotheses are put forward to explain the summer generation pattern: virgin oviposition, and strong intersexual competition.
6. No differences in clutch size were found between the two generations.     

Influence of egg size differences within egg clutches on larval parameters in nine libellulid species (Odonata)

Abstract.  1. In libellulids, egg size differs between species and populations. There are also size differences within egg clutches of individual females.
2. Past experiments suggest that there are two different types of egg clutches in libellulids. Egg size decreases significantly during oviposition in species that perform non-contact guarding during oviposition. In contrast, in species ovipositing in tandem, egg size is randomly distributed.
3. This study deals with the possible consequences of egg size variation within the different egg clutch types. The study examined whether there is a correlation between egg development time, offspring sex or larval size and egg size.
4. The current experiments were conducted in Namibia and Germany. Five non-contact guarding and four tandem guarding libellulid species were used.
5. In some species larger eggs needed more time to develop, in some species no correlation between egg size and egg development time could be found, whereas in other species larger eggs developed faster.
6. The sex ratio was biased towards females in Leucorrhinia dubia and in Sympetrum striolatum and egg size was not associated with gender.
7. In both egg clutch types larger eggs resulted in larger larvae. In this study, evidence was found that the effects of egg size diminished with progressing larval development under good conditions. However, it is possible that the effects may have a greater influence under harsh circumstances.     

Effects of colony size on larval performance in a processionary moth

1. Some lepidopteran species have larvae that live gregariously, especially in early instars. Colony‐living species may benefit from improved protection from predators, thermoregulation, and feeding facilitation, for example. 2. While many studies have compared solitary and gregarious life styles, few data exist as to the relationship between size of the larval colony and larval performance in gregarious species. The present study was aimed at understanding the importance of colony size for growth and survival of the northern pine processionary moth (Thaumetopoea pinivora) larvae. 3. Field studies, comparing three different sizes of colonies of T. pinivora larvae, showed that individuals in larger colonies had a higher survival rate compared with those living in smaller colonies and also a faster growth rate. 4. The higher survival rate of large colonies was attributed to improved protection from predacious arthropods. 5. In early spring, the young larvae bask in the sun to increase their body temperature. In field experiments the thermal gain was higher in large colonies, and individuals in such colonies also grew faster. As growth rate was not affected by colony size when the ability to bask was experimentally removed in a laboratory experiment, the higher growth rate of the larger colonies was probably due to improved thermoregulation rather than feeding facilitation. 6. The size of larval colonies of gregarious insects depends on natural mortality events as well as on female oviposition strategy. Our results show that decreasing colony size can lead to a reduction in growth rate and survival. It is therefore important to understand whether or not small colonies will benefit equally from the gregarious behaviour.     

Superparasitism in Cotesia glomerata: response of hosts and consequences for parasitoids

Abstract.  1. Superparasitism occurs in Cotesia glomerata (Hymenoptera: Braconidae), a gregarious endoparasitoid of Pieris spp. (Lepidoptera: Pieridae). The response of P. brassicae larvae to superparasitism and the consequences for the parasitoid were examined in order to elucidate the ecological significance of this behaviour.
2. Field surveys of a Swiss population revealed that C. glomerata brood sizes from P. brassicae larvae ranged from three to 158, and both the female ratio and the body weight of emergent wasps correlated negatively with brood size. In the laboratory, single oviposition on P. brassicae larvae did not produce any brood size larger than 62, but brood size increased with superparasitism.
3. Laboratory experiments demonstrated that both naive and experienced female wasps were willing to attack hosts that had been newly parasitised by themselves or conspecifics. Superparasitism reduced survivorship but increased food consumption and weight growth in P. brassicae larvae. Superparasitism lengthened parasitoid development and prolonged the feeding period of host larvae.
4. Despite a trade-off between maximising brood size and optimising the fitness of individual offspring, two or three ovipositions on P. brassicae larvae resulted in a greater dry female mass than did a single oviposition on the host. Thus, superparasitism might be of adaptive significance under certain circumstances, especially when host density is low and unparasitised hosts are rare in a habitat.     

Food competition among Scotophugu stercoraria Iarvae with emphasis on its effects on reproductive success

Abstract. 1. Laboratory experiments with yellow dung fly larvae, Scatophaga sfercoraria L. were carried out in the hope of clarifying the causal relationship underlying the extreme variation in body size in wild populations.
2. First, parents were collected in the field and newly hatched larvae exposed to different density levels under two temperature regimes (15°C and 20°C). Second, heritability experiments were done.
3. The results showed that density during the larval stage has a very significant effect on adult size. Male size is more affected than female size and the male larvae spend more time feeding than the female larvae with the result that the males are on average larger. Also, the additive genetic component is of little or no significance, supporting the notion that body size is positively related to fitness.
4. Finally, the results are discussed in relation to field data on the effects of size on both male and female reproductive success.     

The effects of group size, leaf size, and density on the performance of a leaf-mining moth

1. The effects of two factors, leaf size and group size, on the performance of the Tupelo leafminer, Antispila nysaefoliella (Lepidoptera: Heliozelidae), were examined by fitting growth models to mine expansion data using nonlinear mixed-effects models. 2. The rate of mine expansion served as a proxy for larval performance because of its correlation with both feeding activity and growth rate and is also the means by which a larva achieves its final mine size (or total consumption). 3. Leaf size was used as a measure of resource availability, and was expected to reduce the impact of resource competition and enhance larval performance. 4. In contrast to the unidirectional effects expected for leaf size (i.e. more resources should enhance performance), the direction for the effects of group size was expected to depend on the mechanism(s) driving the effect. For example, if there is resource competition among larvae in a group, then this could increase the feeding rates of some larvae or reduce the total consumption of others. However, if leaf mining induces host plant chemical defences, then larger groups might elicit a greater defensive response by the host plant (at the leaf), and hence, be characterized by reduced feeding and growth rates. 5. To investigate these interactions, two growth models, the Gompertz model and a modified version of the von Bertalanffy growth equation, were fitted to time series of the sizes of individual leaf mines using nonlinear mixed-effects models. Linear and nonlinear associations of each factor (group size or leaf size) with model parameters were then evaluated using a hierarchical testing procedure by determining: (i) whether inclusion of the factor produced a better-fit model, and (ii) if it did, the form of that relationship (i.e. linear or nonlinear). 6. Three patterns were detected with these analyses. (i) Leaf size had a significant positive, linear relationship with mine expansion rate. (ii) Group size had a significant quadratic relationship with mine expansion rate. (iii) The effects of leaf and group size on the maximum mine size were opposite to those found with growth rate.     

Effect of nutritional condition on larval food requisition behavior in a subterranean termite Reticulitermes speratus (Isoptera: Rhinotermitidae)

Although optimal investment theory would be similarly applicable to eusocial insects to maximize colony reproductive outputs, directly distinguishing an amount of investment in each larva should be a difficult task for workers because of the characteristics of group living. Thus, it is expected that workers adjust brood care by using a cue or signal conveying information of larval status. In termites, which are typical group of eusocial insects, there are nevertheless few direct observations on worker brood care and little is known about cues inducing worker feeding. I show here that a Japanese subterranean termite Reticulitermes speratus uses an overt food solicitation by larva, “pecking”, as a cue for worker feeding. Direct observations demonstrated that workers feed larvae in response to larval pecking. Furthermore, nutritional experiments showed that larvae exhibited pecking more frequently when their nutrient status is lower; hence, pecking may be an honest reflection of larval hunger status. These results indicate that workers can feed more starved larvae than less starved ones because pecking honestly reflects larval hunger state. That is, feeding in response to pecking should standardize the total amount of food intake of each larva and help a termite colony make worker investment efficient.     

Intraspecific competition amongst larval Aedes aegypti: food exploitation or chemical interference?

Abstract. 1. Competition within and between the larval instars of the yellow fever mosquito, Aedes aegypti , can be measured by its effect on stage duration. In a series of laboratory experiments the relative importance of chemical interference and food exploitation in mediating competition between first and fourth instar larvae was investigated.
2. In contrast to the results of three previous studies, I found no evidence that a chemical growth retardant played any part in greatly prolonging the stage durations of larvae in both age classes.
3. When competition between the two age classes became important, the relative increase in stage duration wasgreater for first instar larvae than for fourth instar larvae. This result can be reproduced with Gilpin & McClelland's (1979) model of competition by food exploitation alone, providing the range of available food particle size is assumed to be an increasing function of age. An additional, but less important, refinement makes the conversion efficiency of food into larval biomass a decreasing function of age.     

Host tree architecture mediates the effect of predators on herbivore survival

Abstract.  1. Vegetation structural complexity is an important factor influencing ecological interactions between different trophic levels. In order to investigate relationships between the architecture of trees, the presence of arthropod predators, and survival and parasitism of the autumnal moth Epirrita autumnata Borkhausen, two sets of experiments were conducted.
2. In one experiment, the architectural complexity of mountain birch was manipulated to separate the effects of plant structure and age. In the other experiment the trees were left intact, but chosen to represent varying degrees of natural complexity. Young autumnal moth larvae were placed on the trees and their survival was monitored during the larval period.
3. The larvae survived longer in more complex trees if predation by ants was prevented with a glue ring, whereas in control trees smaller canopy size improved survival times in one experiment. The density of ants observed in the trees was not affected by canopy size but spider density was higher on smaller trees. The effect of canopy structure on larval parasitism was weak; larger canopy size decreased parasitism only in one year. Until the fourth instar the larvae travelled shorter distances in trees with reduced branchiness than in trees with reduced foliage or control treatments. Canopy structure manipulation by pruning did not alter the quality of leaves as food for larvae.
4. The effect of canopy structure on herbivore survival may depend on natural enemy abundance and foraging strategy. In complex canopies herbivores are probably better able to escape predation by ambushing spiders but not by actively searching ants.     

Impact of balsam fir flowering on pollen and foliage biochemistry in relation to spruce budworm growth, development and food utilization

The impact of balsam fir (Abies balsamea (L.) Miller) flowering on nutritional and allelochemical quality of pollen, current-year and one-year-old foliage is studied in relation to spruce budworm (Choristoneura fumiferana Clem.) (Lepidoptera: Tortricidae) growth, development and utilization of food and nitrogen. In the laboratory, using fresh food from the field, we simulated conditions of low larval population density, in which there is no current-year foliage depletion during the spruce budworm feeding period. Similarly, we simulated conditions of high larval population density when current-year foliage depletion occurs.Because of the high nutritive value of pollen (high amounts of amino acids and minerals, especially nitrogen; low monoterpene content), insects from flowering trees reached the fifth instar five days earlier than those from non-flowering trees, and had heavier dry- and nitrogen-weights at the beginning of the fifth instar. At budbreak, switching from pollen to current-year foliage negatively affected conversion efficiencies and digestibilities of food and nitrogen (AD; ADN; ECDN; ECI; ECIN). The switch from pollen to new foliage had a detrimental impact on fifth-instar survival and on newly-moulted sixth-instar dry- and nitrogen-weights. Moreover, during the fifth instar, balsam fir flowering reduced the nutritive value of current-year foliage, which in turn, might have contributed to the reduced larval growth. Nevertheless, during the sixth instar, balsam fir flowering affected the biochemistry of current-year foliage in ways that enabled larvae to compensate for their low fifth-instar biological performance; larvae also managed to reach pupal dry weight similar to larvae reared on non-flowering trees. Current-year foliage from flowering trees contained less nitrogen, total soluble sugars and total monoterpenes. Those foliar characteristics enabled larvae to increase food and nitrogen consumption rates (RCR; RNCR), because of lower repellency and/or post-ingestional feedback from monoterpenes.As for current-year foliage, balsam fir flowering reduced nitrogen, total soluble sugar and total monoterpene contents in one-year-old foliage during the sixth-instar feeding period. These characteristics enabled sixth-instar larvae, fed on old foliage from flowering trees, to have high relative food and nitrogen consumption rates (RCR; RNCR). Larvae were also able to reach higher relative growth rates (RGR) and relative nitrogen accumulation rates (RNAR) compared to larvae reared on one-year-old foliage from non-flowering trees. Finally, larvae on flowering trees had pupal dry weight similar to those from non-flowering trees, but reached the adult stage nine days earlier.Regardless the foliage type consumed by spruce budworm larvae during the sixth instar, pollen consumption during early larval stages reduced total development time, and thus exposure time to natural enemies. This phenomenon might increase larval survival. Balsam fir flowering changed the biochemistry of one-year-old and current-year foliages, but did not affect pupal dry weights of larvae reared on flowering trees compared to those reared on non-flowering trees. Thus, at low population density, spruce budworm populations in balsam fir flowering stands might be favoured over those in balsam fir non-flowering stands. In addition, when larvae consumed one-year-old foliage during the entire sixth instar, those on flowering trees are probably favoured over those on non-flowering trees. However, because flowering trees produce less new foliage than non-flowering trees, current-year foliage depletion may occur earlier on flowering trees than on non-flowering trees. Thus, at similar larval population density, larvae on flowering trees might have to feed on one-year-old foliage earlier than those on non-flowering trees. In that case, spruce budworm populations on non-flowering stands would be favoured over those on flowering stands.