Are polyphagous geometrid moths with flightless females adapted to budburst phenology of local host species?

The majority of studies demonstrating local adaptation of insect herbivores involve sessile species, particularly those with a parthenogentic phase to their life history or endophagous "parasites" of plants. Current arguments suggest the strength of selection determines whether local adaptation can or cannot take place. Therefore local adaptation should not be limited to species with such traits. We studied the ability of three polyphagous geometrid moths with flightless adult females ( Erannis defoliaria , Operophtera brumata and O. fagata ) to synchronise their egg hatching with the budburst of a local host species in north east Scotland. A strong selection for hatching time is expected among generalist moths given the large variation in budburst phenology and an inability to hatch in synchrony with budburst decreases moth fitness substantially. In two successive seasons, we trapped emerging females from patches of five host species and recorded the temperature sum needed for 50% egg hatch of each brood laid by the trapped females. The hatching times of broods were compared against the average budburst time of the maternal host species in the study area. In addition, the trapping dates of each female were recorded. Only O. brumata showed synchrony with egg hatch and budburst which suggests local phenological adaptation to different host species . This could be maintained by selection and partial reproductive isolation between populations dwelling on different host species. No phenological adaptation was found in the other common geometrids of the study area.     

Growth and development of a generalist insect herbivore, Operophtera brumata, on original and alternative host plants

A generalist feeding strategy is common among eruptive insect herbivores but the ultimate reasons for a generalist strategy are not clear. Although generalist insect herbivores are able to complete their life cycle on several species of host plants, there is wide variation in the performance of individuals grown on different hosts. We examined whether different populations of Operophtera brumata are adapted to use the host species which is locally most abundant, and how the host plant affects growth and development of the insect. We reared two allopatric populations (eastern Finland, Prunus padus; south-west Finland, Quercus robur) on four species of host plants (Pr. padus, Populus tremula, Q. robur, Salix phylicifolia) from neonate larvae to the adult stage and measured the growth and development of individuals and the timing of adult hatching. The performance of both populations was best on Pr. padus, and the south-western population, originally on Q. robur, was well adapted to this host. The host affected the growth of females more than that of males. The host plant had an unexpected effect on hatching times of the adults. Individuals grown on the original host hatched in normal synchrony, i.e. males 6–7 days before females; but on alternative hosts this synchrony was disturbed. As is common in eruptive, capital-breeding generalist moths where female fecundity is linked to weight, host quality is critical for the flightless females of O. brumata. We suggest that in a heterogeneous environment the disturbing effect of alternative host plants on adult emergence may decrease the population density and growth rate compared to the potential maximum in a homogeneous environment. Received: 8 July 1999 / Accepted: 29 October 1999     

Genetic differentiation as a result of adaptation to the phenologies of individual host trees in the galling aphid Kaltenbachiella japonica

Abstract.

• 1 The adaptation of the non-migratory galling aphid Kaltenbachiella japonica (Matsumura) to the budburst phenologies of individual host trees (Ulmus davidiana var. japonica Nakai) was investigated. There was a large variation in budburst time between individual host trees. We tested a hypothesis that aphid populations on respective host trees are genetically differentiated through adaptation to the budburst phenologies of host trees.
• 2 There was significant correlation between the budburst time of eight host trees in two successive years.
• 3 Tree-associated populations differed significantly in egg-hatching time, and the mean hatching time was significantly correlated with the mean budburst time of respective host trees.
• 4 Mating was made between male and female sexuales of the same clone (i.e. self-mating) to obtain selfed eggs. Hatching patterns of selfed eggs showed that there was a substantial amount of genetic variance in hatching time between clones (galls). Nested ANOVA demonstrated that the between-tree component contributed more to the phenotypic variance than the within-tree or within-gall components.
• 5 Reciprocal crosses between males and females of different clones furthermore demonstrated significant differences in hatching time between cross combinations, but no significant differences between the two reciprocals within combinations. This result suggests that there are no maternal effects for the timing of egg hatch and that the differences between selfed lines are attributable to genetic variance.
• 6 The fine-scale adaptation hypothesis was supported by egg-hatching experiments, which further suggest that budburst phenology varying between individual trees is acting as intense selective pressure on the egg-hatching time of associated insects.

     

Effects of photoperiod and raised winter temperatures on egg development and timing of oviposition in the willow psyllid Cacopsylla moscovita

Nymphal development of Cacopsylla moscovita (And.) (Homoptera: Psylloidea) takes place only on female catkins of Salix repens L. and close phenological synchrony is crucial because development times of catkins and nymphs are similar. Eggs are laid on catkins as soon as they develop and close synchrony between oviposition and budburst maximizes time available for nymphal development. Sampling adult C. moscovita in the field revealed little synchrony between egg development and budburst, with over 60% of females containing mature eggs four weeks before catkins first appeared. In the laboratory, egg development was influenced by both temperature and photoperiod. At 10°C, egg development occurred approximately one month earlier than at 5°C and two weeks earlier than in the field. Adult survival in the laboratory was substantially reduced at increased temperature, with only 20% of adults surviving longer than two weeks at 10°C, compared with over 95% at 5°C. Body condition (weight corrected for size) of males and females decreased significantly in the field over winter. However, body condition of females levelled off before budburst, coincident with egg development implying that females had resumed feeding. We discuss our results in relation to regulation of phenological synchrony between oviposition and catkin appearance.     

The influence of plant species on attraction and host acceptance inCotesia glomerata (Hymenoptera: Braconidae)

Females of the larval parasitoidCotesia glomerata (L.) use plant-associated cues to locate their lepidopteran host,Pieris rapae L. In this study we investigated the influence of four host plant species,Brassica oleracea var.acephala (‘Vates’ kale),Tropaeolum majus (nasturtium),Lunaria annua (honesty), andCleome spinosa (spider flower), on two components of the host selection process inC. glomerata, namely, attraction and host acceptance. Choice tests in a flight tunnel showed that parasitoids were attracted to some host plant species more than to others in the absence of host larvae.B. oleracea was the most attractive plant species, followed byL. annua, T. majus, andC. spinosa. In previous studies it was shown thatB. oleracea carries highly suitable hosts forC. glomerata and that, in the field, parasitization rates on this plant were the highest. When host larvae were reared on the four host plant species and then transferred to a common substrate (B. oleracea var.capitata, cabbage), plant species that had served as diet for the hosts did not have a significant effect on acceptance for parasitization. Thus, parasitoids were attracted to host plant species differentially, but they did not discriminate among host larvae based on the dietary history of their hosts. ForC. glomerata, it appears that phytochemistry mediates host selection more by influencing parasitoid attraction than it does by affecting host acceptance.     

Behavior of newly hatched potato tuber moth larvae,Phthorimaea operculella Zell. (Lepidoptera: Gelechiidae), in relation to their host plants

A study was made of neonate larvae of Phthorimaea operculella.Host finding from soil-laid eggs and dispersal from hosts and nonhosts were first examined. Of first-instar larvae hatching from soil-laid eggs, 80% found the potato plant while roughly 50% found each of the other three plants (datura, tobacco, and tomato). Dispersal from potato, datura, and tobacco was very low, while on tomato it was higher and a high mortality was observed in the 24 h period. Dispersal from nonhost plants was high. Behavior on leaves of hosts and non-hosts is described and some host-plant specific behaviors are identified. Effects on behavior of some of the physical and chemical factors are described. Methylene dichloride extracts of potato leaf wax reduced locomotion rates and the number of turns during locomotion. Also methylene dichloride extract and, to a lesser extent, methanol extract caused biting behavior.     

Diet-induced food preference by Manduca sexta larvae: acceptable non-host plants elicit a stronger induction than host plants

Diet-induced changes in food preference by fifth instar larvae of the tobacco hornworm, Manduca sexta (Johan.) (Lepidoptera, Sphingidae), were examined. Two groups of larvae with different diet experience were used: larvae reared on a host or on an acceptable non-host plant species. Each group of larvae was offered a choice between leaf discs from each rearing plant species (2-plant choice test) and food preference was measured as the consumption of one plant species relative to that of the other plant species. Diet-induced changes in preferences were larger with the host versus acceptable non-host plant pairs Solanum pseudocapsicum (L.) versus Raphanus sativus (L.), Lycopersicon esculentum (Mill.) versus Vigna sinensis (Savi), and Datura innoxia (L.) versus V. sinensis than with the host versus host plant pairs L. esculentum versus Capsicum annuum (L.) and L. esculentum versus D. innoxia. To examine how much the food preference had been altered for each test plant species alone, two other groups of larvae were offered a choice between leaf discs from a single plant species and filter paper discs laced with distilled water (1-plant choice test). Larvae preferred the dietary plant species more strongly than the non-dietary plant species in tests using the following plant species: for C. annuum with C. annuum and L. esculentum as diets, for V. sinensis with V. sinensis and L. esculentum or D. innoxia as diets, and for R. sativus with R. sativus and S. pseudocapsicum as diets. The preference for the hosts L. esculentum and D. innoxia did not change significantly after rearing larvae on different hosts or on an acceptable non-host. Thus, diet-induction by M. sexta larvae results in an enhancement of preference for the dietary plant species which is much stronger with acceptable non-hosts than with hosts.     

Teratocytes of the solitary endoparasitoid Meteorus gyrator (Hymenoptera: Braconidae): morphology, numbers and possible functions

Abstract. Laboratory studies investigated the development of teratocytes derived from the eggs of the parasitoid Meteous gyrator (Thun.) in its host, the tomato moth Lacanobia oleracea (L.). At hatching, each parasitoid egg produced an average of approximately 1000 teratocytes, but this number declined to approximately 400 during the course of parasitism. The teratocytes increased in size markedly, such that 7 days after egg hatch their mean diameter was approximately four times that of the cells immediately after dissociation. The haemolymph of parasitized hosts had reduced phenoloxidase activity, and teratocytes inhibited phenoloxidase activity when coincubated with plasma from nonparasitized hosts. The injection of teratocytes into nonparasitized fifth‐instar L. oleracea larvae suppressed growth and induced a supernumerary moult in some larvae. A number of parasitism‐specific proteins were detected in the haemolymph of parasitized hosts, and incubation of teratocytes in culture media indicated that these cells were a source of at least two of these proteins.     

Predicted changes in the synchrony of larval emergence and budburst under climatic warming

Summary The impact of climatic warming on the synchrony of insect and plant phenologies was modelled in the case of winter moth (Operophtera brumata) and Sitka spruce (Picea sitchensis) in the Scottish uplands. The emergence of winter moth larvae was predicted with a thermal time requirement model and the budburst of Sitka spruce was predicted from a previously published model (Cannell and Smith 1983) based on winter chilling and thermal time. The date of emergence of winter moth larvae was predicted to occur earlier under climatic warming but the date of budburst of Sitka spruce was not greatly changed, resulting in decreased synchrony between larval emergence and budburst. The general question of how a change of climate might affect phenological synchrony and insect abundance is discussed.     

Performance of leaf beetle larvae on sympatric host and non-host plants

Studies asking the ability of insects to utilize novel host plants often use novel hosts that are allopatric with the insect population under investigation. However, since the outcomes of species interactions are often site-specific, such studies cannot tell us whether a plant would actually be used by a given insect population if the plant grew sympatrically with it. We therefore performed a quantitative genetics experiment to analyse the performance of larvae of the leaf beetle Oreina elongata Suffrian (Coleoptera: Chrysomelidae, Chrysomelinae) on two host and three non-host plants, collected from a site where insects and plants co-occur in the Western Alps. When raised on the non-host Petasites albus (L.), larvae were able to survive equally well as on the two hosts, Adenostyles alliariae (Gouan) and Cirsium spinosissimum (L.), whereas they did not survive on the two other non-hosts, Peucedanum ostruthium (L.) and Rumex alpinus L. On P. albus, growth rate was slightly lower and development time slightly longer than on the two hosts. We found a genotype by environment interaction only for growth rate but not for development time and survival. However, the shape of the reaction norms of growth rates suggests that it is unlikely that selection could favour the inclusion of P. albus into the host range of the study population.     

Parasitism by Pteromalus cerealellae (Hymenoptera: Pteromalidae) on the Cowpea weevil, Callosbruchus maculatus (Coleoptera: Bruchidae): Host density, temperature effects, and host finding ability

Studies on interactions between a larval parasitoid, Pteromalus cerealellae (Boucek) and one of its hosts, Callosobruchus maculatus (F.) were carried out in the laboratory. The number of host larvae parasitized by P. cerealellae increased with host larvae at low densities and tended to a plateau at a density of 25 larvae per female parasitoid. Each parasitoid was able to parasitize more hosts and produced more offspring at 20 and 25 °C than at 30 °C. The number of non-infested seeds mixed with seeds infested with the last instar of C. maculatus did not preclude P. cerealellae from identifying infested seeds and attacking larvae inside them. When infested seeds were tightly packed, several host larvae escaped parasitism. P. cerealellae may be a useful biological control agent in newly harvested cowpea with low C. maculatus infestation, and lowering the temperature of the storage system may enhance the effectiveness of this parasitoid.     

Fecundity and Longevity of Green Vegetable Bug,Nezara viridula, Following Parasitism byTrichopoda giacomellii

The effects of parasitism by the ArgentinianTrichopoda giacomellii(Blanchard) on reproduction and longevity of its host,Nezara viridula(L.) are reported. Parasitoid larvae suppress egg maturation, reducing by 70% the fecundity of mature female hosts during the period of larval development. Egg viability was not affected, but mating frequency was reduced by approximately 50%. When parasitized as newly eclosed adults, 84% of females fail to reproduce. In male hosts, fertility and mating frequency were not affected during the period of larval parasitoid development. In male and reproductively immature female hosts, death was coincident with, or occurred shortly after parasitoid emergence (2–4 days); in mature females, death occurred on average 2 weeks after larval parasitoid emergence. Host mortality occurred as a consequence of tissue damage incurred as the parasitoid larvae emerged from the host. Some individuals survived parasitism though no further reproductive activity (mating or oviposition) occurred. The effectiveness ofT. giacomelliias a biological control agent is discussed in relation to its impact on reproduction and survival of its host and contrasted with the action of otherTrichopodaspecies.     

Varying degree of physiological integration among host instars and their endoparasitoid affects stress‐induced mortality

In natural populations of insect herbivores, genetic differentiation is likely to occur due to variation in host plant utilization and selection by the local community of organisms with which they interact. In parasitoids, engaging in intimate associations with their host during immature development, local variation may exist in host quality for parasitoid development. We compared the development of a gregarious endoparasitoid, Cotesia glomerata L. (Hymenoptera: Braconidae), collected in The Netherlands, in three strains and three caterpillar instars (L1–L3) of its main host, Pieris brassicae L. (Lepidoptera: Pieridae). Hosts had been collected in The Netherlands and France, and were reared in the laboratory for one generation. We also used an established Dutch laboratory strain that had not been exposed to parasitoids for at least 24 generations. Parasitoid survival to adulthood was inversely correlated with host instar at parasitism. Adult parasitoid body mass was largest when hosts were parasitized as L1 and smallest when hosts were parasitized as L3, whereas egg‐to‐adult development time was quickest on L3 hosts and slowest on L1 hosts. Higher survival and faster development of C. glomerata on French L2 hosts also showed that there is variation in host‐instar‐related suitability. Many L2 and most L3 caterpillars that were parasitized exhibited signs of pathogen infection and perished within a few days of parasitism, whereas this never happened when hosts were parasitized as L1 or in non‐parasitized control caterpillars. Our results reveal that, irrespective of the host strain, L1 hosts are optimally synchronized with C. glomerata development. By contrast, the high precocious mortality of L3 larvae may be due to stress‐induced regulation by the parasitoid in order to ‘force’ its developmental program into synchrony with the developing parasitoid larvae. Our results underscore a potentially important role played by pathogens in mediating herbivore–parasitoid interactions that are host‐instar‐dependent in their expression.     

Lack of physiological improvement in performance of Callosamia promethea larvae on local host plant favorites

Summary As a species, the promethea silkmoth, Callosamia promethea (Saturniidae: Lepidoptera) exhibits a wide host range on 6–10 families of plants, although specific populations are known to have local foodplant favorites. We tested the hypothesis that larvae from a particular host plant lineage would show physiological adaptations to this host compared with larvae from other host plant lineages. We found no evidence that larval survival and growth was any better for larvae fed the natural plant of the parental population than for larvae from other host lineages. These natural host lineages include: black cherry (Prunus serotina Ehrh.), tuliptree (Liriodendron tulipifera L.), sassafras (Sassafras albidum (Nutt.) Nees) and spicebush (Lindera benzoin (L.) Blume). The only apparent manifestation of physiological specialization was the inability of tuliptree lineages of C. promethea to survive on paper birch (Betula papyrifera Marsh), although this may reflect the geographical pattern of adaptation to birch, rather than a negative correlation with adaptation to tuliptree. These results suggest that for C. promethea larvae, growth performance and survival is primarily influenced by plant nutritional quality, rather than physiological adaptations to the locally preferred host plant.     

The effect of host age ofLymantria dispar larvae [Lep.: Lymantriidae] on the development ofGlyptapanteles liparidis [Hym.: Braconidae]

The endoparasitic development ofG. liparidis was examined in 3 different host stages of gypsy moth larvae. Hatching ofG. liparidis-larvae occurred 3 to 5 days after oviposition in hosts parasitized during their premoulting period, and after 5 to 7 days in those parasitized in the 3^rd midinstar state. The parasites generally moulted to the 2^nd larval instar between the 11^th and 13^th day in the first group, and between the 13^th and 15^th day in the latter, when they had reached a volume of 0.04–0.05 mm³. The positive correlation between host ecdysis and the ecdysis of 1^st stadium larvae to L₂ suggested that host moulting influenced the development of the parasitoid larvae. Emergence from the host larvae occurred at 20°C after 27 days on average, and coincided with the parasites moulting to the 3^rd instar. Five to 7 days after spinning their cocoons near the developmentally arrested host larva, the male, and 1 to 2 days later the female wasps eclosed. Due to the variation in the number of parasites per host, no difference was observed between the hosts parasitized at various stages; however, a tendency for later parasitized hosts to contain more parasite larvae was evident. The nutritional conditions of the moth parental generation influenced both host and parasite development. On the other hand no influence of host age was observed on emergence dates of larvae and wasps.      

Host selection and utilisation of Sonchus oleraceus (Asteraceae) by Helicoverpa armigera (Lepidoptera: Noctuidae): A genetic analysis

The cotton bollworm (Helicoverpa armigera) prefers the common sowthistle (Sonchus oleraceus L.) to cotton (Gossypium hirsutum L.), sorghum (Sorghum bicolor L.) and maize (Zea mays L.) for oviposition in the field in Australia. Using the common sowthistle and cotton as host plants, we carried out this study to evaluate genetic variation in both oviposition preference and larval growth and genetic correlation between maternal preference and larval performance. There was a significant genetic component of phenotypic variation in both characters, and the heritability of oviposition preference was estimated as 0.602. Helicoverpa armigera larvae survived slightly better and grew significantly faster on common sowthistle than on cotton, but genetic correlation between maternal preference and larval growth performance was not detectable. Instead, larval growth performance on the two hosts changed with families, which renders the interaction between family and host plant significant. As a result, the genetic correlation between mean values of larval growth across the two host species was not different from zero. These results are discussed in the context of the relationship between H. armigera and the common sowthistle and the polyphagous behaviour of this insect in general.     

Asynchrony in larval development of the pine beauty moth, Panolis flammea, on an introduced host plant may affect parasitoid efficacy

In the United Kingdom, Panolis flammea (Den. and Schiff.) (Lepidoptera: Noctuidae) is an important pest species of the introduced lodgepole pine but not of its natural host Scots pine. The timing of P. flammea larval growth must be synchronized with its host tree if the larvae are to succeed. We collected field data during 1990 which revealed that the phenological window starts earlier in Scots pine and is shorter than that observed in lodgepole pine. The larvae are found in the field earlier and within a narrower time frame within a Scots pine forest than in a lodgepole pine forest. The larval developmental period is significantly longer on lodgepole pine than on Scots pine. The synchrony/asynchrony of P. flammea to its natural host (Scots pine) and an introduced tree (lodgepole pine) results in the parasitoids having a different impact on the larvae of the two hosts. At any one time, the host plant, caterpillars and parasitoids are more synchronous on the ancestral Scots pine than on lodgepole pine, resulting in a higher percentage of larvae in the optimal instar for parasitism at that time. In lodgepole pine, the percentage of suitable instars available to parasitoids is lower at any given time. The information presented here furthers our understanding of the possible mechanisms for the observed differential population dynamics of the insect on Scots pine and lodgepole pine in the UK. Handling editor: Robert Glinwood.     

Larval dispersal from potential hosts within a population of a generalist herbivore, Choristoneura rosaceana

The dispersal behavior of Choristoneura rosaceana (Harris) (Lepidoptera: Tortricidae) first instar larvae was studied in the laboratory. The objectives were to investigate the proximal factors influencing larval dispersal and to establish whether a correspondence exists between larval host acceptance and performance. A dispersal bioassay was validated by demonstrating the presence of a positive correlation between larval host acceptance in the laboratory and in the field. Larval age and family origin, as well as host species attributes were shown to influence larval dispersal rates. Seasonal changes in host plants slightly changed the rank order of larval host acceptance. Leaf texture and the availability of refuges on host plants seemed to be important factors influencing the rate of larval dispersal. Plant odor appeared to be used by the larvae to locate leaves. Nitrogen content of plant species corresponded to larval dispersal rates, but the cause of this association is unclear. Larval dispersal did not match host suitabilities as measured by larval performance. The relationship between host preference and suitability in the obliquebanded leafroller is discussed in an ecological and evolutionary perspective.     

Effects of different Beauveria bassiana isolates on field populations of Lygus lineolaris in pigweed (Amaranthus spp.)

The tarnished plant bug, Lygus lineolaris (Palisot de Beauvois), is a pest of various fruit, vegetable, fiber, and seed crops; including cotton. Lygus spp. populations often build on alternate host plants before moving to cotton, and in the midsouthern U.S. wild host plants, such as pigweed (Amaranthus spp.), play a major role in L. lineolaris population development. Three isolates of the entomopathogenic fungus Beauveria bassiana (Balsamo) were evaluated for L. lineolaris control in redroot pigweed (Amaranthus retroflexus L.): one from L. lineolaris in Mississippi (TPB3); one from Lygus hesperus (Knight) in California (WTPB2); and one commercial isolate from Mycotrol^® (GHA). Fungal applications resulted in moderate to high mycosis in adults (33 to 80%) and moderate mycosis in nymphs (36 to 53%) that were collected from field plots at 2 days post-treatment and incubated under laboratory conditions. Although TPB3 was previously found to be more pathogenic in laboratory bioassays, there was not a consistent separation of this isolate from the other two isolates in field trials. Where differences in adult mycosis or mortality were observed, TPB3 was the most pathogenic. However, in one field trial 7 day mortality for nymphs treated with GHA was higher than those treated with TPB3 or WTPB2. Infection rates at 2, 7, and 14 days post-treatment from caged and non-caged adults suggested that movement of adults among plots occurred, which could have masked some treatment effects. Fungal treatments did not significantly reduce populations relative to controls. This may have been caused by delayed mortality rates under field conditions and/or difficulties with estimating population change under field conditions characteristic of wild host plant populations (e.g., heterogeneous populations, adult movement, and small plot size). Further work evaluating time–dose–mortality over dynamic temperatures, spring and fall field trials on this and other wild hosts, and improved methods for estimating populations on wild hosts are needed.     

Allorhogas pyralophagus [Hym.: Braconidae], a gregarious external parasite imported into texas,USA, for biological control of the stalkborerEoreuma loftini [Lep.: Pyralidae] on sugar cane

Allorhogas pyralophagus Marsh is a recently discovered doryctine braconid with potential for field trials against stalk borers infesting both large- and small- stemmed Graminacea. This external gregarious parasite was originally reared fromEoreuma loftini (Dyar) larvae infesting Johnson grass [Sorghum halepense (L.) Pers.] and sugar cane (Saccharum officinarum L.) from the states of Sinaloa and Nuevo Leon, Mexico. The parasite is easily reared in the laboratory using a wide range of host larvae [Diatraea spp.,E. loftini, Chilo sp. andTryporyza nivelli (F)]. Hosts are placed in grass stems or paper straws for exposure to ovipositing females.A. pyralophagus has been field released and recovered from the following hosts infesting sugar cane:E. loftini andD. saccharalis (F.) in Texas.Diatraea spp. in Trinidad,D. rufescens in Bolivia andT. nivella in Sumatra, Indonesia. Approved as Contribution No. TA-20743 of the Texas Agricultural Experiment Station.