Performance of larval parasitoid,Bracon brevicornis on two Spodoptera hosts: implication in bio-control of Spodoptera frugiperda

Successful pest management using parasitoids requires careful evaluation of host-parasitoid interactions. Here, we report the performance of larval ecto-parasitoid wasp, Bracon brevicornis (Wesmael) on important agricultural pests, Spodoptera litura (Fabricius) and S. frugiperda (J.E. Smith). Biology of B. brevicornis was studied on different host instars under laboratory and cage setup. In no-choice assay, the parasitoid development was highest on fifth-instar S. litura larvae as the wasp laid?~?253 eggs with 62% hatching, 76% pupae formation and 78% adult emergence. Similarly, these parameters were highest on fifth instar S. frugiperda larvae (293 eggs, 57% hatching, 80% pupae formation, 70% adult emergence). In two-choice assay, B. brevicornis preferred fourth or fifth over third instar larvae of both hosts. Successful parasitism depends on host paralysis and suppression of host immunity. B. brevicornis interaction downregulated cellular immunity of both hosts as shown by reduced hemocyte viability and spreading. The percent parasitism rate of B. brevicornis was unaltered in the presence of host plant, Zea mays in cage study. 76 and 84% parasitism was observed on fifth-instar larvae of S. litura and S. frugiperda, respectively. We evaluated the performance of B. brevicornis as a biocontrol agent on S. frugiperda in maize field. Our results show 54% average reduction in infestation after release of B. brevicornis. Taken together, we report the performance of B. brevicornis on important insect pests for the first time in laboratory and field conditions. Our findings indicate that B. brevicornis is a promising candidate for integrated pest management.

     

Competitive interactions of a new invader (Spodoptera frugiperda) and indigenous species (Ostrinia furnacalis) on maize in China

Spodoptera frugiperda has been reported in China since the beginning of 2019, threatening maize production and raising questions about interactions with indigenous pests on maize. Spodoptera frugiperda is in the same feeding guild as the indigenous species Ostrinia furnacalis, with both pests attacking maize plant tissues such as whorls and tassels. In this study, the interaction of these species was assessed under laboratory and field conditions by examining the survival and predation rate of larvae in maize whorls and tassels. Intraspecific and interspecific interactions of larvae were assessed to characterize movements in arenas in the presence and absence of food. When S. frugiperda and O. furnacalis were present at an equal ratio, S. frugiperda prevailed over O. furnacalis with a high survival rate (over 90%) and predation rate of O. furnacalis (over 40%) under most scenarios in both laboratory and field conditions. When both species occurred at different ratios in the field, S. frugiperda had on average almost 16 times the number of survivors as O. furnacalis on whorls, and 8.3 times the number on tassels, highlighting a substantial advantage in these environments. Spodoptera frugiperda larvae undertook a low frequency of lethal attacks (strikes) and showed a higher incidence of defensive movements compared to O. furnacalis. A field survey showed that S. frugiperda larvae can colonize vegetative plant stages in maize, but it remains to be seen if competitive interactions can be linked to population dynamics of both species though they are expected to favor S. frugiperda.

     

Attack and defense movements involved in the interaction of <Emphasis Type="Italic">Spodoptera frugiperda</Emphasis> and <Emphasis Type="Italic">Helicoverpa zea</Emphasis> (Lepidoptera: Noctuidae)

The fall armyworm, Spodoptera frugiperda (J. E. Smith) and the corn earworm, Helicoverpa zea (Boddie) are among the main pests of maize. Both species exhibit cannibalistic behavior and quite often share the same feeding guild in maize (maize ears), which can result in several interspecific and intraspecific interactions. Paired interaction scenarios of intraspecific and interspecific larvae were assessed in arenas in the presence and absence of food to characterize movements resulting from interactions of these insects. There was a difference in the frequency of behavioral movements in all the interactions, except for S. frugiperda in the presence of food. Head touching and recoiling were the predominant movements in most of the interaction scenarios. Spodoptera frugiperda exhibited a predominance of defensive movements when competing against H. zea in the same instars. Cannibalism and predation occurred frequently in interactions involving 6th instar of H. zea against opponents in 4th instar. Larvae of H. zea show a higher aggressive movement than S. frugiperda. The larvae of S. frugiperda take advantage during the interactions, although they present more defensive movements compared to H. zea. This study provides relevant information regarding the interaction of these species and intraguild interaction, which might influence the population dynamics and the competitive displacement of pest species that share the same ecological niche.     

Antifeedant activity and toxicity of leaf extracts from <Emphasis Type="Italic">Porteresia coarctata</Emphasis> Takeoka and their effects on the physiology of <Emphasis Type="Italic">Spodoptera litura</Emphasis> (F.)

Antifeeding and toxic effects of chemical extracts from Porteresia coarctata Takeoka on the insect pest Spodoptera litura (F.) were investigated in the laboratory. Hexane extracts of P. coarctata leaves were re-dissolved in dimethyl sulfoxide and applied in water to leaves of Ricinus communis L. (castor). A concentration of 1,000 and 2,000 ppm extract exhibited significant antifeeding and toxic effects to third instar larvae when compared to the same concentration of dimethyl sulfoxide in water as control. One day after feeding on extract-treated castor leaves, the mortality of S. litura larvae was 50%. Mortality increased to 95% after 11 days at a dosage of 2,000 ppm. Compared to the control, third instar larvae fed on castor leaves treated with P. coarctata leaf extract at different concentrations showed significant reduction in protein and DNA content in the fat body and midgut tissues. Results demonstrate the insecticidal and antifeedant properties of P. coarctata leaf extract against the generalist pest S. litura leading to biochemical changes in the insect body. Thus, P. coarctata contains phytochemicals, and the extract, in crude and pure form, has potential as a botanical insecticide in alternative control strategies against Lepidoptera pests.     

The effects of 2,4-dihydroxy-7-methoxy-1,4-benzoxazin-3-one on two species of <Emphasis Type="Italic">Spodoptera</Emphasis> and the growth of <Emphasis Type="Italic">Setosphaeria turcica</Emphasis> in vitro

Maize seedlings contain high amounts of glucosidically bound 2,4-dihydroxy-7-methoxy-1,4-benzoxazin-3-one (DIMBOA). The effects of DIMBOA on the feeding behaviour and performance of two noctuids, Spodoptera exigua Hübner and S. frugiperda Smith, were compared. The question was raised whether S. frugiperda, preferring maize and other Poaceae, is better adapted to DIMBOA than S. exigua. In addition, the effects of DIMBOA on the mycelial growth of the plant pathogen Setosphaeria turcica Leonard et Suggs (causal agent of northern corn leaf blight) was assessed in vitro. DIMBOA had an antifeedant effect on S. exigua but stimulated feeding in S. frugiperda in dual-choice experiments. In a no-choice setup, larvae of S. exigua gained less biomass and had a prolonged development when feeding on an artificial diet containing DIMBOA. However, pupal weight was not significantly different between treatments. In contrast, larvae of S. frugiperda were not affected by DIMBOA. Strong detrimental effects of DIMBOA were found on the mycelial growth of the pathogen S. turcica.     

Semiochemical and natural product-based approaches to control Spodoptera spp. (Lepidoptera: Noctuidae)

The genus Spodoptera contains some of the most destructive crop pests in the world owing to the worldwide distribution of the different species and their wide host ranges. Uses of semiochemicals for insect control have been recently reviewed, but strategies developed specifically against Spodoptera species are scarce. In this review, we present an updated account of the semiochemicals and other natural product-based approaches to monitor and control the most damaging Spodoptera spp. In general, successful control of Spodoptera littoralis, S. frugiperda, S. litura, and S. exigua have been reported through mass trapping, mating disruption, and attract-and-kill methods. An updated survey of the pheromone components of these species and an outlook of future perspectives against these pests are also outlined.     

Bioinsecticidal effect of the flavonoids pinocembrin and quercetin against <Emphasis Type="Italic">Spodoptera frugiperda</Emphasis>

Flavonoids function in many aspects of plant–insect interactions, but the responses of insects to these compounds vary greatly. In this study, we determined the effects of two widely distributed flavonoids, pinocembrin and quercetin, on the feeding behavior, survival, and development of the fall armyworm Spodoptera frugiperda J.E. (Smith) (Lepidoptera: Noctuidae). In a choice test, S. frugiperda larvae strongly rejected leaves treated with pinocembrin at concentrations of 10, 50, or 100 μg/cm². Larvae fed normally on leaves treated with quercetin at 10 and 50 μg/cm², but showed 57 % deterrence when fed on leaves treated with 100 μg/cm² quercetin. At concentrations of 0.01–1 µg/cm², pinocembrin and quercetin functioned as phagostimulants for S. frugiperda. In a multiple-choice experiment, S. frugiperda larvae preferred to consume untreated leaves or those treated with 0.1 µg/cm² pinocembrin, but rejected leaves treated with 5–50 µg/cm² pinocembrin. In a no-choice feeding experiment, larvae fed on leaves treated with 5 and 50 μg/cm² pinocembrin consumed less than those fed on leaves treated with 0.1 and 1 μg/cm² pinocembrin or untreated leaves. Pinocembrin at 1–50 μg/cm² negatively affected larval weight and survival, thus showing a toxic effect. In contrast, leaf consumption and larval weight were not significantly affected by quercetin at 0.1, 1, 5, and 50 μg/cm², and mortality rates only slightly increased. Because of its dual activity, pinocembrin could be used for insect control in a stimulo-deterrent diversionary strategy: the same compound could promote both stimulate (low doses) and deter insect activity (high doses).     

Bioactivities of caffeic acid methyl ester (methyl-(E)-3-(3,4-dihydroxyphenyl)prop-2-enoate): a hydroxycinnamic acid derivative from Solanum melongena L. fruits

Feeding deterrent and growth inhibitory effects of caffeic acid methyl ester (CME) isolated from the fruit extract of eggplant, Solanum melongena L. along with two known phenolic acids, caffeic acid (CA) and chlorogenic acid (CG) and azadirachtin (taken as active control), were evaluated against Spodoptera litura Fab. and Achaea janata L. (Lep., Noctuidae). The structure of CME was determined on the basis of extensive spectroscopic (UV, FTIR, ¹H, and ¹³C NMR) data analysis. CME has exhibited strong feeding deterrent activity against S. litura and A. janata larvae with ED₅₀ values of 52.02 and 41.02 μg/cm² leaf area, respectively. CA and CG were relatively less effective compounds when compared with CME. The results also revealed that the isolated compound CME caused marked larval growth inhibition in S. litura and A. janata larvae after 7 days of feeding on a treated diet. The effect on growth of larvae was concomitant with the reduced feeding and digestibility of ingested food. So the activities of digestive proteases within the midgut of larvae were examined using specific substrates to understand the digestive physiology in the larval guts. The bulk of the activity was associated with serine proteases comprising trypsin-, chymotrypsin-, and elastase-like enzymes, which decreased in diets containing CME, whereas, interestingly increased with CG. From the results, we conclude that CME isolated from S. melongena fruit extract shows feeding deterrent and larval development inhibitory activities that may prove useful in the management of S. litura and A. janata larvae.     

Antifeedant and toxic effects of acetogenins from <Emphasis Type="Italic">Annona montana</Emphasis> on <Emphasis Type="Italic">Spodoptera frugiperda</Emphasis>

Several species of the large family of tropical plants Annonaceae have been intensely investigated over the last 20 years, mainly because of the discovery of annonaceous acetogenins. These compounds are powerful cytotoxics, with potential applications as insecticides, antiparasitics, acaricides, fungicides, and antitumor drugs. Annona montana Macfad. (Annonaceae) grows in Santa Cruz de la Sierra, Bolivia, where an infusion of leaves is used for the treatment of lice, influenza, and insomnia. The major acetogenins from a Bolivian collection of A. montana, annonacin (1), cis-annonacin-10-one (2), densicomacin-1 (3), gigantetronenin (4), murihexocin-B (5), and tucupentol (6), were evaluated for their antifeedant and toxic effects on Spodoptera frugiperda Smith (Lepidoptera: Noctuidae), a serious pest affecting corn crops in Argentina. All the acetogenins produced 100% mortality during the larval or pupal stages at 100 μg of treatment per gram of diet. In addition, compounds 2, 3, and 4 deterred more than 80% feeding at the same dose. Relative toxicity values of LD₅₀ for the strongest larvicidal compounds 1, 2, and 4 were determined, indicating that the three compounds are effective natural insecticides. This is the first report on the antifeedant and toxic effects produced by the particular type of acetogenins, the mono-THF acetogenins, on the lepidopteran S. frugiperda. No correlation was detected between the toxicity of the mentioned compounds to larvae and the known capacity of the acetogenins 1, 2, and 4 to inhibit the NADH oxidase, indicating that the inhibition of the mitochondrial complex I is not the only cause for larval mortality of S. frugiperda.     

Field evolved resistance to carbamates,organophosphates, pyrethroids,and new chemistry insecticides in <Emphasis Type="Italic">Spodoptera litura</Emphasis> Fab. (Lepidoptera: Noctuidae)

Spodoptera litura Fab. (Lepidoptera, Noctuidae) is a polyphagous pest and has been shown to be resistant to a wide range of insecticides, which has led to sporadic out breaks of the pest and failure of crops. We were interested to establish if resistance to insecticides is diverse in different populations of S. litura collected from various areas with variable temperatures. We collected S. litura from eight different locations and compare the toxicity of insecticides. Resistance to the pyrethroids ranged from 20- to 11,700-fold compared with the Lab-PK (laboratory susceptible population). The resistance to bifenthrin was the lowest in a population collected from Multan and the highest to esfenvalerate in a population collected from Lodhran. Similarly, very high level of resistance to spinosad, indoxacarb, and methoxyfenozide was observed in all eight populations. In contrast, resistance to organophosphates was significantly lower than the pyrethroids, spinosad, indoxacarb, and methoxyfenozide, while toxicity of emamectin to field populations was similar to the Lab-Pk. The results are discussed in relation to integrated pest management (IPM) for S. litura with special reference to management of field evolved resistance to insecticides.     

Zur Wirkung der Nahrungspflanze auf die Entwicklung und Vermeharung der Baumwolleule,Spodoptera littoralis (Boisd.) (Lep.,Noctuidae)

Effect of some host plants on development and reproduction of Spodoptera littoralis (Boisd.) (Lep., Noctuidae) Investigations were performed to study the effect of some hostplants on development and reproduction capacity of the Egyptian cotton leaf-worm,Spodoptera littoralis (Boisd,), Larval and pupal weights, life span, average number of eggs laid per female and percent digestability for fifth instar larvae were measured. Cotton leafworm was able to complete its life cycle only when reared on eigher castor bean leaves, clover, sweet potato leaves, or the outer green cabbage leaves. Chlorophyl-free inner white cabbage leaves, maize, and grape vine are nutritionally insufficient to support growth and development of larvae and, in turn, cause increase in span of survived larvae and none of these larvae metamorph to pupae. It seems that there is a positive correlation between larval and pupal weights and average number of eggs laid per female; the greater the weight of larvae and pupae the higher the number of eggs laid. Castor bean leaves seem to be the most favorable host plant for rearing cotton leafworm in the laboratory. The differences and variations in the growth and development of cotton leafworm due to feeding on different host plants are discussed.     

Antifeedent and toxic activity of endophytic Alternaria alternata against tobacco caterpillar Spodoptera litura

Fungal endophytes play a significant role in plant resistance to insects by altering the nutritional quality of plants or by production of various alkaloid-based defensive compounds. Ethyl acetate extract of endophytic Alternaria alternata isolated from Azadirachta indica A. Juss was tested for its bioactivity against Spodoptera litura (Fab.). Effect of different concentrations of ethyl acetate extract of A. alternata was assessed on growth and development of S. litura larvae. The results indicate higher larval mortality on diet supplemented with the fungal extract than control. Adverse effects of A. alternata were also observed on development of the insect. Ethyl acetate extract of the fungus also significantly reduced the adult emergence, longevity and reproductive potential of S. litura. Larvae feeding on diet supplemented with fungal extract showed significant reduction in relative growth and consumption rate, as well as efficiency of ingested and digested food. The antifeedant and toxic effects of A. alternata may be due to production of bioactive molecules by this fungus.     

Characterization of priming,induced resistance,and tolerance to Spodoptera frugiperda by silicon fertilization in maize genotypes

Fall armyworm (FAW) Spodoptera frugiperda is considered the main defoliating insect pest of maize in many countries. Silicon (Si) applied to plants has been shown to increase the resistance to insects, especially in grasses such as maize. This study characterized the effects of Si fertilization regarding priming, induced resistance, and tolerance to FAW in a landrace variety and hybrid of maize. Si was applied in soil of potted-plants as H₄SiO₄ at 2 t ha^?1 when maize plants were at V2 stage, and when they reached V3 two FAW neonates were placed in the plant’s whorls to cause herbivory. FAW performance was evaluated on excised leaf sections in the laboratory and on plants with larvae infesting V4-stage plants in the greenhouse. Concentrations of H₂O₂, malondialdehyde, and Si, and the activities of antioxidant enzymes were recorded and correlated to Si-based responses on plant growth, and FAW injury and performance. As main results, there was reduced FAW injury and larval weight gain in Si-treated plants subjected to herbivory. Greater root dry mass was observed in the landrace variety with Si and without herbivory. Landrace plants showed higher shoot weights than the hybrid under FAW infestation. Si-fertilized plants showed higher H₂O₂ concentrations. The highest peroxidase activities occurred in Si-treated plants without herbivory, and the catalase and superoxide dismutase activities were highest in Si-treated plants without herbivory or herbivory-injured plants without Si. In conclusion, Si-based defense in maize to FAW involve mixed effects of priming and tolerance, and were more pronounced in the landrace variety.

     

Parasitism byMeteorus rubens onAgrotis ipsilon as affected by supplementary food and kairomone, field studies

Field experiments in clover field showed that releasing of 10 matedM. rubens females/m², increased parasitism amongA. ipsilon larvae from 16.7% at 0 to 43,3% after 10 days. Also, releasing 10 females/m² after spraying 3% molasses in tomato field increased parasitism from 4% at 0-day to 27.5% after 10 days amongA. ipsilon larvae. Spraying molasses and kairomone together on maize plants, increased markedly the rate of parasitism amongS. cretica larvae comparing with parasitism obtained when each of them was sprayed alone. The release ofM. rubens in a rate of 10 females/m² after spraying molasses and kairomone induced a significant increase in the rate of parasitism amongS. cretica larvae after 10 and 15 days compared to releasing 3 or 4 females of the parasitoid. A significant increase in parasitism amongS. cretica larvae on tomato after 10 and 15 days was recorded whenM. rubens was released at the rates of 5 and 10 females/m², after spraying molasses and kairomone, compared to other treatments.     

Wirkungen einesBacillus thuringiensis-Präparats und eines Juvenilhormon-Analogons über den Erdboden aufSpodoptera littoralis (Boisd.) (Lep.,Noctuidae)

Summary Sensitivy of Spodoptera littoralis (Boisd.) to soils treated with preparations of Bacillus thuringiensis and of a juvenil hormone analogue Various instar larvae ofSpodoptera littoralis were exposed to soil treated, with aBacillus thuringiensis-containing preparation (Bactospeine) and a juvenile hormone analogue (Ro 20–36 000).After treatment with Bactospeine, only high concentrations could affect emergence. Suppressed egg laying and sterility were the most marked effects. Also, the produced eggs were smaller in size.The juvenile hormone analogue showed high activity, particularly against freshly moulted 6th. instar larvae. Greater effects were elicited after ecposing the larvae to treated soil than to treated leaves for 48 hours. It appears to be fairly stable in soil. Its morphogenetic activity could be detected 60 days after application.     

Castilleja and Pedicularis confirmed as telial hosts for Cronartium ribicola in whitebark pine ecosystems of Oregon and Washington

The primary objective of this research was to determine whether native species of Castilleja and Pedicularis are naturally infected by Cronartium ribicola in whitebark pine ecosystems of the Oregon and Washington Cascade Range, USA. Secondary objectives were to monitor the phenology of aecial and telial hosts to determine whether there is sufficient time for C. ribicola to complete its life cycle within high‐elevation stands and to evaluate the variety of susceptible native host species within these genera through field and growth chamber inoculation. These objectives were approached through fieldwork in 2008 and 2009 in whitebark pine ecosystems at Mt. Rainier, Mt. Adams, Mt. Hood, Mt. Bachelor, Tumalo Mtn. and Crater Lake. Forty‐nine observational study plots were established and monitored. Natural C. ribicola infection was detected on 84 Pedicularis racemosa plants and five Castilleja plants (C. applegatei, C. miniata and C. parviflora). Field observation provided evidence that there is sufficient time for C. ribicola to complete its life cycle on hosts within high‐elevation whitebark pine stands. In 2009, 18 field inoculation plots were established at Mt. Rainier and Crater Lake. Field inoculation confirmed the susceptibility of two additional species within these genera, C. arachnoidea and P. bracteosa. All four Castilleja species inoculated in the growth chamber developed infection, with an overall infection incidence of 62% (167 out of 270 plants). The identity of the rust species on field specimens as C. ribicola was verified through PCR and sequencing of the ITS1‐5.8S‐ITS2 region of DNA. Improved understanding of the role of these newly recognized hosts in white pine blister rust epidemiology should be used to prioritize sites for the restoration of ecologically valuable whitebark pine.     

Infectivity prospects of both nematodes and bacterial symbionts against cotton leafworm, Spodoptera littoralis (Biosduval) (Lepidoptera: Noctuidae)

A study was made on the susceptibility of the cotton leafworm, Spodoptera littoralis (Biosd.), to the nematode, Steinernema carpocapsae (All) and Heterorhabditis bacteriophora. Three concentrations of each nematode species were used (75, 150, and 300 infective juveniles) for each treatment. The nematode, H. bacteriophora, gave 100% mortality 96, 90, and 48 h, respectively, post-treatment of S. littoralis larvae with 75, 150, and 300 infective juveniles. On the other hand, S. carpocapsae (All) gave 100% mortality 120, 90, and 56 h post-treatment, respectively. Therefore, H. bacteriophora was more potent against cotton leafworm than S. carpocapsae. Studies on the infestation intensity of the nematode species against the cotton leafworm showed the predominance of H. bacteriophora over S. carpocapsae, while in studies of the reproductive rate those of S. carpocapsae predominate. The bacterial symbionts of each nematode species were isolated and tested alone against cotton leafworm at concentrations ranging between 1×10² and 5×10³ bacterial cells/larva. The results showed a higher activity of Photorhabdus luminescens than that of Xenorhabdu nematophilus.     

Activities of digestive and detoxification enzymes in multiple generations of beet armyworm, <Emphasis Type="Italic">Spodoptera exigua</Emphasis> (Hübner), in response to transgenic Bt cotton

To reveal the multi-generation effects of transgenic Bt cotton on the metabolism of secondary target herbivores, the activities of digestive and detoxification enzymes in beet armyworm, Spodoptera exigua (Hübner), fed with transgenic Bt (cv. GK-12) vs. non-Bt cotton (cv. Simian-3) for three successive generations were examined. Significantly lower activities of lipase, carboxylesterase (CarE) and acetylcholinesterase (AChE), and higher activities of trypsin and total superoxide dismutase (T-SOD) were observed in Bt-fed S. exigua than in non-Bt fed individuals during the three generations. But the activity of amylase did not differ between the Bt-fed and non-Bt fed S. exigua. Among the three generations of Bt-fed S. exigua, the activities of lipase and trypsin decreased significantly in the second or third generation than in the first generation. But those of CarE and AChE were increased significantly. Among the three generations of non-Bt fed S. exigua, the activity of lipase was significantly higher in the second and third generations than in the first generation. The increasing adaptation and fitness of S. exigua after continuous exposure to Bt cotton was supposed to be associated with the enhanced activities of detoxification enzymes. Measuring multi-generational enzymes activities of S. exigua in response to Bt cotton can provide proofs of its adaptation development, which is suggested to be a mandatory part of risk assessment of transgenic Bt plants on secondary target herbivores.     

Among-tree variation in leaf traits and herbivore attacks in a deciduous oak,Quercus dentata

Abstract

Variations in defensive and some other leaf traits were studied in a population of an oak species, Quercus dentata Thunberg, in northern Japan, with reference to attacks by ectophagous herbivores and leafminers. The oak population showed substantial individual variations in concentrations of total phenolics and condensed tannins, nitrogen content, trichome density, leaf area and budburst timing. With the exception of leaf mass per area, which showed a positive relation with leaf toughness and a negative relation with water content, no significant relation was observed between the plant traits studied, suggesting an absence of trade-off or linkage between them. The oaks also showed substantial individual variations in leaf area loss by ectophagous herbivores, densities of major leafminers (Phyllonorycter and Stigmella species) and survival of Phyllonorycter sap-feeding larvae. The density of trichomes showed a significant, negative relation with leaf area loss by ectophagous herbivores, but significant, positive relations with densities of some leafminers. The other leaf traits seldom showed significant relations with herbivore densities or survival. In this oak population, these traits may not have enough variations to be reflected in the abundance and performance of herbivores.