Structural plasticity of olfactory neuropils in relation to insect diapause

1. Many insects that live in temperate zones spend the cold season in a state of dormancy, referred to as diapause. As the insect must rely on resources that were gathered before entering diapause, keeping a low metabolic rate is of utmost importance. Organs that are metabolically expensive to maintain, such as the brain, can therefore become a liability to survival if they are too large.
2. Insects that go through diapause as adults generally do so before entering the season of reproduction. This order of events introduces a conflict between maintaining low metabolism during dormancy and emerging afterward with highly developed sensory systems that improve fitness during the mating season.
3. We investigated the timing of when investments into the olfactory system are made by measuring the volumes of primary and secondary olfactory neuropils in the brain as they fluctuate in size throughout the extended diapause life‐period of adult Polygonia c‐album butterflies.
4. Relative volumes of both olfactory neuropils increase significantly during early adult development, indicating the importance of olfaction to this species, but still remain considerably smaller than those of nondiapausing conspecifics. However, despite butterflies being kept under the same conditions as before the dormancy, their olfactory neuropil volumes decreased significantly during the postdormancy period.
5. The opposing directions of change in relative neuropil volumes before and after diapause dormancy indicate that the investment strategies governing structural plasticity during the two life stages could be functionally distinct. As butterflies were kept in stimulus‐poor conditions, we find it likely that investments into these brain regions rely on experience‐expectant processes before diapause and experience‐dependent processes after diapause conditions are broken.
6. As the shift in investment strategies coincides with a hard shift from premating season to mating season, we argue that these developmental characteristics could be adaptations that mitigate the trade‐off between dormancy survival and reproductive fitness.

     

The impact of experimental fire regimes on seed production in two tropical eucalypt species in northern Australia

Abstract This study investigated the effect of three experimental fire regimes on the fecundity, ovule development and seedfall of two common wet-dry tropical savanna eucalypts, Eucalyptus minima and Eucalyptus tetrodonta, in northern Australia. Both species flower early in the dry season and ovule development occurs during the dry season. This coincides with a period of frequent fires. The three fire regimes considered were applied for four years between 1990 and 1994. These regimes were (i) Unburnt, (ii) Early, fires lit early in the dry season, and (iii) Late, fires lit late in the dry season. The treatments were applied to nine catchments (15–20 km²) with each fire regime replicated three times. Fire intensity typically increases as the dry season proceeds. Therefore, early dry season fires generally differ from late dry season fires in both their intensity and their timing in relation to the reproductive phenology of the eucalypts. Late dry season burning significantly reduced the fecundity of both species, whereas Early burning had no significant effect. Ovule success was significantly reduced by the Early burning for both species. The Late burning significantly reduced ovule success in E. tetrodonta, but not in E. miniata. The results suggest that fire intensity and fire timing may both be important determinants of seed supply. Fire intensity may be a determinant of fecundity, whereas fire timing in relation to the reproduction phenology may have a significant impact on ovule survival. Both fire regimes resulted in a substantial reduction in seed supply compared with the Unburnt treatment. This may have a significant impact on seedling regeneration of these tropical savanna eucalypts.     

Reproductive Phenology of Woody Species in a North Australian Tropical Savanna1

Interspecific and interannual variation in reproductive phenology was quantified for 50 common species of trees and shrubs from a mesic savanna near Darwin, northern Australia. The presence of buds, flowers, and fruit was noted over a 30-month period, from September 1992 to February 1995. Surveys were undertaken at monthly intervals for the less common species, and at bimonthly intervals for ten of the common trees and tall shrubs. The majority of species flowered each year at about the same time. There was no evidence of sub-annual or continuous regimes of reproductive phenology. There was no supra-annual carryover of seed-bearing fruit in the canopy of any species. The peak flowering periods were the mid to late dry season (July–August) and the transition between the dry season and the wet season (October–November). The two dominant trees–Eucalyptus miniata and E, tetrodonta– flowered during the dry season, thereby providing resources for some elements of the vertebrate fauna. Flowering and fruiting were uncommon at the end of the wet season (February/March), although two species that flower and fruit at this time (E. porrecta and Terminalia ferdinandianas may provide resources to consumers at a time when floral or fruit resources are otherwise scarce. Because the peak of reproductive activity takes place during the late dry season, fruit maturity and seed dispersal have occurred prior to the onset of the rainy season for most species, and germination and seedling establishment potentially may take effect in response to the first rains. Late dry season fires, which tend to be extensive and intense, are a potential threat to the floral and fruit reserves within these savannas.     

Asymmetric life-history decision-making in butterfly larvae

In temperate environments, insects appearing in several generations in the growth season typically have to decide during the larval period whether to develop into adulthood, or to postpone adult emergence until next season by entering a species-specific diapause stage. This decision is typically guided by environmental cues experienced during development. An early decision makes it possible to adjust growth rate, which would allow the growing larva to respond to time stress involved in direct development, whereas a last-minute decision would instead allow the larva to use up-to-date information about which developmental pathway is the most favourable under the current circumstances. We study the timing of the larval pathway decision-making between entering pupal winter diapause and direct development in three distantly related butterflies (Pieris napi, Araschnia levana and Pararge aegeria). We pinpoint the timing of the larval diapause decision by transferring larvae from first to last instars from long daylength (inducing direct development) to short daylength conditions (inducing diapause), and vice versa. Results show that the pathway decision is typically made in the late instars in all three species, and that the ability to switch developmental pathway late in juvenile life is conditional; larvae more freely switched from diapause to direct development than in the opposite direction. We contend that this asymmetry is influenced by the additional physiological preparations needed to survive the long and cold winter period, and that the reluctance to make a late decision to enter diapause has the potential to be a general trait among temperate insects.     

Species traits help explain butterfly habitat use in a tropical forest

1. When species can access all parts of the landscape, species-sorting metacommunity theory predicts that community composition depends on habitat choice and interactions with other species and the environment. These filtering processes can also depend on species' traits.
2. The authors investigated how traits mediate a species-sorting process in determining butterfly community composition in a naturally patchy landscape in the tropical Western Ghats, India. The authors asked, do traits mediate access to certain habitats and does seasonality affect these patterns? The authors surveyed 56 habitat patches in three habitat types: laterite plateau grasslands, ridge grassland, and moist-deciduous forest, in a 65-km² landscape.
3. Non-palatable butterflies showed similar occurrences across seasons and habitats, but palatable butterflies were less commonly encountered in open habitats in the dry season. Polyphagous butterflies occurred infrequently in the dry season in laterite habitats, potentially indicating emigration or diapause patterns are linked to diet breadth.
4. All species were present in all habitats, implying dispersal does not limit access to different habitat patches, consistent with the species-sorting metacommunity concept. Nevertheless, butterfly occurrence was strongly influenced by the interaction of mobility and habitat type with sedentary species occurring less often in low-resource open laterite patches than mobile species.
5. Species sorting is typically regarded as occurring directly through environmental filters, but here the authors suggest that the environmental filter acts through movement limitations. Studies integrating landscape heterogeneity and species characteristics will help us better understand metacommunities and species distributions in nature.

     

Effect of fire regime on plant abundance in a tropical eucalypt savanna of north‐eastern Australia

Abstract Changes in plant abundance within a eucalypt savanna of north‐eastern Australia were studied using a manipulative fire experiment. Three fire regimes were compared between 1997 and 2001: (i) control, savanna burnt in the mid‐dry season (July) 1997 only; (ii) early burnt, savanna burnt in the mid‐dry season 1997 and early dry season (May) 1999; and (iii) late burnt, savanna burnt in the mid‐dry season 1997 and late dry season (October) 1999. Five annual surveys of permanent plots detected stability in the abundance of most species, irrespective of fire regime. However, a significant increase in the abundance of several subshrubs, ephemeral and twining perennial forbs, and grasses occurred in the first year after fire, particularly after late dry season fires. The abundance of these species declined toward prefire levels in the second year after fire. The dominant grass Heteropogon triticeus significantly declined in abundance with fire intervals of 4 years. The density of trees (>2 m tall) significantly increased in the absence of fire for 4 years, because of the growth of saplings; and the basal area of the dominant tree Corymbia clarksoniana significantly increased over the 5‐year study, irrespective of fire regime. Conservation management of these savannas will need to balance the role of regular fires in maintaining the diversity of herbaceous species with the requirement of fire intervals of at least 4‐years for allowing the growth of saplings >2 m in height. Whereas late dry season fires may cause some tree mortality, the use of occasional late fires may help maintain sustainable populations of many grasses and forbs.     

Caryedon serratus and its parasitoids in the Savanna around Lamto,Ivory Coast

In the savanna around Lamto,Caryedon serratus develops, in the beginning of the dry season, on maturing seeds ofPiliostigma thonningii (Caesalpinioideae). A 2^nd generation, and sometimes a 3^nd one, develops in dry seeds. There is no pupal diapause during the rainy season and adults seems to spend this period in a semi-lethargic state. The parasitoid complex ofC. serratus is a chalcid-dominated one. It includes 5 species: one oophagous (Uscana caryedoni) and 4 larvo-nymphal parasitoids. Three species are regular (Anisopteromalus caryedophagus, Bracon sp. andU. caryedoni), the others are sporadic species (Proconura serratocida, Eurytoma caryedocida). Biological data are given for each species.      

Differences in photoperiod‐induced diapause plasticity among different populations of the bark beetle Ips typographus and its predator Thanasimus formicarius

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Environmental Cues,Endocrine Factors,and Reproductive Diapause in Male Insects

Environmental cues, mostly photoperiod and temperature, mediated by effects on the neuroendocrine system, control reproductive diapause in female insects. Arrest of oocyte development characterizes female reproductive diapause, which has two major adaptive functions: It improves chances of survival during unfavorable season(s), and/or it confines oviposition to that period of the year that is optimal for survival of the eggs and progeny. Although reproductive diapause is less well studied in male insects, there may be no sex-dependent differences in regard to the first of these functions. The second one, however, is not valid for the male; instead, selection pressure directs the male's reproductive strategy toward maximum chances of fertilization of the female's eggs with minimum waste of energy. Therefore, in species with female reproductive diapause, the males may or may not exhibit diapause, but if they do, their diapause must be adapted to that existing in conspecific females. Male reproductive diapause is defined as a reversible state of inability of the male to inseminate receptive females. In relation to reproductive diapause, there are several patterns of coadaptations between male reproductive strategy and timing of female receptivity, (a) In some insects, the females are receptive in the early part of their diapause; mating occurs during this period and there is no diapause in the male. The male dies shortly after copulation and the female stores the sperms to fertilize the eggs that develop after termination of the female's diapause, (b) In some species, as in the grasshopper Anacridium aegyptium, females are receptive during diapause; though oocyte development is arrested, copulation occurs and the stored sperms fertilize the eggs when the female's diapause ends. Males were claimed to have no diapause, but recent studies have revealed the presence of a reproductive diapause in a proportion of the males. This and other cases show that female receptivity during reproductive diapause may or may not be accompanied by male reproductive diapause. If there is a reproductive diapause in the male, it is controlled by the same endocrine mechanism, the corpora allata (CA), as in the females, (c) In many species females are refractory during their diapause. In these cases, males exhibit reproductive diapause, which may be light, as in the beetle Oulema melanopus, or well established, as in certain grasshoppers, butterflies, and beetles. In the latter cases, male diapause is controlled by similar environmental cues (photoperiod, temperature) and by the same intrinsic mechanism (neuroendocrine system, especially CA) as female diapause. Nevertheless, male diapause is less intense; the environmental cues leading to its termination are less complex and/or less extreme, so male diapause terminates before that of the females. Presumably, male diapause is under two antagonistic selection pressures: A male should not waste energy by courting dia-pausing refractory females, but he should be ready to copulate as soon as the females become receptive, otherwise he may lose in the competition between males for females. Some further strategies, which do not seem to fit the above patterns, are also outlined.     

Phenotypic plasticity, seasonal climate and the population biology of Bicyclus butterflies (Satyridae) in Malawi

Abstract.

• 1 Seasonal polyphenism is studied in a community of five African butterflies of the genus Bicyclus at the transition between a wet and a dry season from May to July.
• 2 Butterflies characterized by large eyespots and, especially in B.sufitza (Hewitson), a pale band (the wet season form) are replaced over this period by butterflies lacking conspicuous wing markings (the dry season form, dsf). The latter butterflies also tend to be larger, but more variable in size. Butterflies of an intermediate phenotype are recruited over a comparatively short interim period.
• 3 This turnover coincides with a period of declining temperature and drying of the habitat, including the grasses on which larvae feed. Butterflies are progressively more likely to rest on brown leaf litter rather than on green herbage.
• 4 A relationship with temperature is supported by laboratory experiments with B.saJitza and B.anynana (Butler) showing that increasingly extreme dsf butterflies develop with decreasing rearing temperature in the final larval instar.
• 5 Some differences in behaviour and activity were observed between the seasonal forms. Butterflies of the dsf develop ovarian dormancy and fat bodies. They can survive to reproduce at the beginning of the rains in November.
• 6 Capture-recapture experiments showed that the adult butterflies have a comparatively long life expectancy and are quite sedentary.
• 7 The results are discussed in relation to a hypothesis linking the polyphenism to seasonal changes in resting background and selection for crypsis.

     

Seasonal polyphenism in the wild: survey of wing patterns in five species of Bicyclus butterflies in Malawi

Abstract.

• 1 For three successive years Bicyclus butterflies were caught on a daily basis at a field site in Malawi.
• 2 Over 5000 butterflies, belonging to five species, were captured. Eight characters describing their plastic wing pattern were measured and analyzed.
• 3 Broadly speaking, the plasticity is similar for all species, with the wet season forms having conspicuous wing markings (e.g. eyespots), and dry season forms lacking these markings.
• 4 However, at a more detailed level, each species has its own specific form of plasticity with especially clear differences in the number of intermediate forms.
• 5 Females generally show a higher degree of plasticity than males.
• 6 The relative frequencies of intermediate forms and the difference between the sexes are associated with differences between the species in their preference for more open habitats or forests.
• 7 The species with the most divergent plasticity is also ecologically and phylogenetically comparatively distant from the others.

     

Soil temperature and depth of legume germination during early and late dry season fires in a tropical eucalypt savanna of north‐eastern Australia

Abstract Temperatures that significantly increase seed germination of some tropical legumes (i.e. 80–100°C) were documented in the topsoil during the passage of early (May) and late (October) dry season fires in a tropical eucalypt savanna of north‐eastern Australia. Elevated temperatures penetrated at least 30 mm into the soil during the higher‐intensity, late dry season fires, but were only detected at 10 mm during the early dry season fires. The depth from which germination of two native legume forbs Galactia tenuiflora and Indigofera hirsuta occurred was positively related to the temperature elevation in the topsoil and was greater after late compared with early dry season fires. A broader range in germination depth, resulting in higher seedling densities, was recorded for I hirsuta after late dry season fires. These results suggest that seedling emergence of native leguminous forbs is likely to occur at a greater density after late rather than early dry season fires in tropical eucalypt savannas of north‐eastern Australia. Therefore, the season of burning, as a result of its relationship to fire intensity, can influence species composition through its effect on seed germination.     

Variation in the life cycle of monogonont rotifers: Commitment to sex and emergence from diapause

1. A review of research on life-cycle events in field and laboratory populations of monogonont rotifers shows that there is great variation at multiple levels: (1) degree of sexual dimorphism; (2) occurrence and timing of sex; (3) propensity for sex during sexual periods; (4) factors controlling initiation of sex; and (5) timing and extent of emergence from diapause. There is no regular pattern where: (1) fertilised resting eggs hatch to start the growing season; (2) populations develop via female parthenogenesis during favourable conditions; and then (3) bisexual reproduction with resting-egg production occurs during later, unfavourable conditions.
2. Sexual reproduction in natural populations can occur throughout much of the growing season, be restricted to some period(s) during the growing season, or be completely absent. During sexual reproduction in both natural and laboratory populations, only some fraction of females produces males or resting eggs. This bet-hedging strategy can prevent a population crash and permits future population growth via female parthenogenesis. Selection against sexual reproduction, and rapid loss of sex, can occur.
3. Laboratory experiments with pond-dwelling species have identified specific environmental factors that induce sex in different species: (1) increasing population density; (2) dietary tocopherol (vitamin E) and (3) long photoperiods. These factors generally are associated with favourable conditions for population growth and production of energy-rich resting eggs: (1) large population size; (2) high probability of contacts between males and fertilisable females; and (3) nutritious diets. Endogenous factors can inhibit responses to these environmental inducers, and thus favour female parthenogenesis.
4. The timing of resting-egg hatching depends on: (1) occurrence of specific environmental conditions; (2) the minimum duration of obligate diapause; and (3) the genotype and physiology of females producing resting eggs. Hatching may occur shortly after oviposition, after a long diapause before or at the start of a new growing season, or throughout the growing season. Hatching can be massive and contribute substantially to population growth and genetic diversity.
5. Areas for future research include: (1) determining the timing and extent of sex and resting-egg hatching in more natural populations, especially those that are marine, benthic, sessile, and interstitial; and (2) identifying environmental and physiological factors controlling these events.

     

Germinable soil seed banks in a tropical savanna: seasonal dynamics and effects of fire.

Abstract The germinable soil seed bank of a tropical eucalypt savanna of north‐eastern Australia was found to be dominated by grasses and forbs, with seed bank density ranging from 58 to 792 seeds per square metre, from a total of 53 species. Late dry season fires and the fire‐related cues, heat shock and smoke, broke the seed dormancy of a range of tropical savanna species. Heat shock promoted the germination of the species groups natives, exotics, subshrubs, ephemeral and twining perennial forbs, and the common species Indigofera hirsuta, Pycnospora lutescens and Triumfetta rhomboidea. Exposure to smoke at ambient temperature promoted germination from the soil seed bank of the species groups combined natives, upright perennial forbs and grasses, as well as the common grasses Digitaria breviglumis and Heteropogon triticeus. The germinable soil seed bank varied seasonally, increasing from the mid wet season (February) and early dry season (May) to a maximum in the late dry season (October). The effect of recent fire history on soil seed bank dynamics was limited to the immediate release of some seed from dormancy; a reduction in seed densities of subshrubs and monocots, other than grasses, in recently burnt savanna; and enhanced seed density of the ephemeral I. hirsuta in the year following fire. The seed banks of most savanna species were replenished in the year following burning.     

Pollination and Breeding System of two Co-Occurring Hirtella Species (Chrysobalanaceae) in Central Brazil

The phenology, major floral characteristics, breeding systems and fruiting success of two co-occurring species of Hirtella: H. glandulosa and H.gracilipes (Chrysobalanaceae) were studied in Central Brazil. The two species occur as trees in mesophyllous forests but H. glandulosa is frequent also in dense savanna areas. Both species flower at the end of the dry season (September) and have flowers with one-day longevity. In both species flowers produce nectar and are pollinated exclusively by butterflies. The number of visits recorded and pollination rate in each species were significantly different and indicate that pollinators prefer H. glandulosa flowers. Natural and controlled fruit sets were low in both species. The index of self-incompatibility (ISI) was 0.17 in H. glandulosa and 0.86 in H. gracilipes. Although ISI in H. glandulosa could denote an incompatibility system, the presence of fruits with aborted embryos at different degrees of development point to an inbreeding depression situation. Low pollen viability and fruit set in H. gracilipes suggested reproductive problems which may be linked to hybridization events.     

Phytoplankton community structure and environment in the Kenyan waters of Lake Victoria

• 1 Phytoplankton species composition, numerical abundance, spatial distribution and total biomass measured as chlorophyll a concentration were studied in relation to environmental factors in September 1994 (dry season) and March 1995 (rainy season), respectively, in the Kenyan waters of Lake Victoria; 103 species were recorded.
• 2 Blue‐green algae (Cyanophyceae) were most diverse, followed by diatoms (Bacillariophyceae), green algae (Chlorophyceae) and dinoflagellates (Dinophyceae).
• 3 Twinspan separated the phytoplankton communities in the Nyanza Gulf and those in the open lake during both seasons. During the dry season, the Nyanza Gulf was strongly dominated by blue‐greens, while diatoms dominated in the open lake. During the rainy season, blue‐greens remained dominant in the Nyanza Gulf although the number of species found was lower than during the dry season; in the open lake, blue‐greens replaced diatoms as the dominant group and there were more species than in the dry season.
• 4 Canonical correspondence analysis indicated that the phytoplankton species distribution was significantly correlated with turbidity during the dry season and with SiO₂ during the rainy season. Chlorophyll a concentrations ranging from 2.0 to 71.5 mg m^‐3 in the dry season and 2.0–17.2 mg m^‐3 in the rainy season confirm earlier reports of increasing phytoplankton biomass in Lake Victoria since the 1960s.

     

Seasonal influence on reproduction in chimpanzees of gombe national park

Although wild chimpanzees are not seasonal breeders, there are seasonal effects on several aspects of chimpanzee reproduction. I examined the seasonal incidence of anogenital swelling in cyclic, pregnant, and acyclic female chimpanzees in Gombe National Park, May 1975–April 1992, and surveyed important reproductive events to determine whether there is a seasonal effect. I analyzed data by season (wet vs. dry) and seasonal quarter;early dry season = May–July;late dry = August–October;early wet = November–January;late wet = February–April. When data for the 17 years are combined, the percentage of females in each reproductive state remains consistent throughout the year. In a given month, 30–35% of subjects were in the cyclic category, 11–15% were pregnant, and 54–61% were acyclic. Cyclic females showed full swelling more often during the late dry season. Pregnant females exhibited anogenital swelling more often during the late dry and early wet seasons. Acyclic females also exhibited a seasonal effect with more anogenital swelling during the late dry season. There is no seasonal difference in frequency of live births (dry, 20;wet, 23). However, the timing of conception showed a seasonal effect (dry, 32;wet, 16). Consistent with earlier reports, the onset of postpartum cycles is highly seasonal;30 occurred during dry season, 9 during wet season. The occurrence of first full swellings for young females is also concentrated in the late dry season. It appears that the dry season is a time of great change for Gombe chimpanzee reproductive physiology. Previous studies indicated that seasonal changes in food availability play a role in increasing group size during the dry season and social contact between females can enhance cyclicity. Accordingly, I suggest that seasonal changes in diet may play a role, either directly (food content) or indirectly (social contact), to alter reproductive physiology.     

Variable chemical defence in the checkerspot butterfly Euphydryas gillettii (Lepidoptera: NymphaIidae)

Abstract.

• 1 Like other checkerspots, Euphydryas gillettii butterflies may contain the defensive chemicals, iridoid glycosides, which are sequestered from their hostplants during larval feeding.
• 2 We analysed the iridoid glycoside content of E.gillettii adults from two different populations, Warm Lake, Idaho, and Granite Creek, Wyoming, that have different patterns of hostplant use.
• 3 Gas chromatographic analysis of thirty butterflies from the Wyoming population showed that they contained a mean of 1.27 (±0.19 SE) % dry weight iridoid glycosides. Notably, 20% of these butterflies contained no detectable iridoid glycosides.
• 4 In contrast, nineteen butterflies from the Idaho population contained a mean of 3.89 (±0.38 SE) % dry weight iridoid glycosides, and all butterflies contained iridoid glycosides.
• 5 These results illustrate how the chemical defence of herbivorous insects varies according to differential use of potential hostplants.

     

The reproductive biology and diapause of the British fungal-breeding Drosophila

Abstract.

• 1 Seasonal reproductive development was investigated in four obligate fungivorous species, D.phalerata Meigen, D.transversa Fallen, D.cameraria Haliday and D.confusa Staeger.
• 2 Samples of females taken from woodlands over a 2 year period were scored for reproductive status and fat content.
• 3 Populations of D.phalerata and D.cameraria maintained in an outdoor insectary provided information on developmental time and adult mortality throughout the year.
• 4 Experiments in controlled environmental chambers gave percentage diapause curves against day length for all four species.
• 5 In D.phalerata the influence of temperature and fungus upon diapause behaviour was also investigated.
• 6 In D.phalerata the optimal strategy for reproductive development can vary from month to month. Maturation of females may be immediate, dependent upon a fungal cue or completely prevented.

     

Seasonal shifts in host usage in Uroleucon gravicorne (Homoptera: Aphididae) and implications for the evolution of host alternation in aphids

ABSTRACT.

• 1 Uroleucon gravicome (Patch) feeds on annual Erigeron species and on perennial Solidago species.
• 2 To test whether Solidago and Erigeron are analogous to winter and summer hosts of typical host-alternating aphids, reproductive performance, host preferences, and host associations were measured for E.strigosus, S.juncea and S.nemoralis during spring, summer and autumn.
• 3 Caged individuals can reproduce on both genera throughout the season, though colonies are rare on Solidago during summer.
• 4 Developmental rate, adult weight, and fecundity decline between May and August on all hosts; however, the drop is least on E.strigosus, intermediate on S.nemoralis, and greatest on S.juncea.
• 5 Throughout the season, reproductive performance is at least as great on Erigeron as on Solidago, the difference being least in spring.
• 6 Soluble nitrogen content shows the same seasonal trends as reproductive performance, declining in all hosts, but declining least in E.strigosus.
• 7 Preference tests and field records show that U.gravicorne moves from Solidago to Erigeron when the first alatae mature in late spring, feeds on Erigeron during the summer, and, in late summer, returns to Solidago where overwintering eggs are laid.
• 8 Solidago is not nutritionally superior during spring or autumn; its use during those seasons may result from the need for good oviposition sites.