Larval host plant affects fitness consequences of egg size variation in the seed beetle Stator limbatus

Egg size variation often has large effects on the fitness of progeny in insects. However, many studies have been unable to detect an advantage of developing from large eggs, suggesting that egg size variation has implications for offspring performance only under adverse conditions, such as during larval competition, periods of starvation, desiccation, or when larvae feed on low-quality resources. We test this hypothesis by examining the consequences of egg size variation for survivorship and development of a seed-feeding insect, Stator limbatus, on both a low-quality (Cercidium floridum) and a high-quality (Acacia greggii) host plant. Our results are consistent with the hypothesis. S. limbatus larval performance was affected by egg size only when developing on the poor-quality host (C. floridum); larvae from large eggs survived better on C. floridum than those from small eggs, while there was no evidence of an effect of egg size on progeny development time, body weight, or survivorship when larvae developed on A. greggii. These results indicate intense selection for large eggs within C. floridum-associated populations, but not in A. greggii-associated populations, so that egg size is predicted to vary among populations associated with different hosts. Our results also support this hypothesis; females from a C. floridum-associated population (Scottsdale) laid larger eggs than females from an A. greggii-associated population (Black Canyon City).     

Host-associated differences in fitness within and between populations of a seed beetle (Bruchidae): effects of plant variability

Summary Experiments were conducted with the sexually reproducing seed beetle Stator limbatus and its hosts in north-central Arizona to determine if it was substructured into units, each specialized for higher fitness on a specific host species. Unlike many studies, we incorporated scale, i.e., conducting experiments between and within beetle populations on seeds from within and between plant species. Of particular interest was whether intraspecific plant variability prevented beetle specialization within beetle populations. Results suggest that S. limbatus is specialized to certain hosts. On the palo verde Cercidium floridum, beetles originally reared from this host had significantly higher emergence compared to beetles transferred from other hosts. We did not test directly for a genetic basis for this. Alternative hypotheses of variation in symbiotic microorganisms in larval guts and maternal effects were assessed. Essentially no bacteria, yeast or protozoa were found, and maternal effects as expressed by varying egg weights were not detected; however, other microorganisms might have been present and maternal effects through inducible enzymes was possible. Caution, then, is needed in any genetic interpretations of our results. The differences on C. floridum were detected from tests between and within beetle populations. Evidence for specialization was not detected on the other hosts, Cercidium microphyllum and Acacia greggii. On the other hosts, beetles performed well regardless of their source. Significant differences were detected among individual plants of C. floridum as to the suitability of their seeds for deveoopment of S. limbatus. No such differences were detected among the other host plants. These patterns of conspecific plant variability are opposite of what is expected if plant variability prevents specialization of beetles to particular species of hosts. Thus, the data suggest seed variability among plants does not prevent specialization to host species in this system. We discuss how the patterns of host use in this study relate to the hypothesis of sympatric host race formation.     

Host-associated fitness variation in a seed beetle (Coleoptera: Bruchidae): evidence for local adaptation to a poor quality host

The geographic distributions of many generalist herbivores differ from those of their host plants, such that they experience coarse-grained spatial variation in natural selection on characters influencing adaptation to host plants. Thus, populations differing in host use are expected to differ in their ability to survive and grow on these host plants. We examine host-associated variation in larval performance (survivorship, development time, and adult body weight) and oviposition preference, within and between two populations ofStator limbatus (Coleoptera: Bruchidae) that differ in the hosts available to them in nature. In one population,Acacia greggii (Fabaceae: Mimosoideae) andCercidium microphyllum (Fabaceae: Caesalpininoideae) are each abundant, while in the second population onlyC. floridum andC. microphyllum are present. In both populations, egg-to-adult survivorship was less than 50% onC. floridum, while survivorship was greater than 90% onA. greggii. Most of the mortality onC. floridum occurred as larvae were burrowing through the seed coat; very low mortality occurred during penetration of the seed coat ofA. greggii. Significant variation was present between populations, and among families (within populations), in survivorship and egg-to-adult development time onC. floridum; beetles restricted toCercidium in nature, without access toC. floridum, survived better and developed faster onC. floridum than beetles that had access toA. greggii. Large host effects on body size were detected for female offspring: females reared onA. greggii were larger than those reared onC. floridum, whereas male offspring wee approximately the same size regardless of rearing host. Trade-offs between performance onC. floridum andC. floridum were not detected in this experiment. Instead, our data indicate that development time and survivorship onC. floridum may be largely independent of development time and survivorship onA. greggii. Patterns of oviposition preference corresponded to the observed patterns of host suitability: in laboratory preference tests, beetles with access toA. greggii in nature tended to prefer this host more than beetles without access to this host in nature.     

Clutch size manipulations in two seed beetles: consequences for progeny fitness

Seed beetles (Coleoptera: Bruchidae) lay their eggs on discrete resource patches, such that competition among larvae for food is an important component of their biology. Most seed beetles, including Stator limbatus, lay eggs singly on individual seeds and avoid superparasitism except when seeds are limiting. In contrast, S. beali, a closely related congener, lays eggs in clutches on a single seed. We tested the hypothesis that natural selection on larval life history characters favors small clutches (selection against large clutches) in S. limbatus, but that selection against large clutches is relaxed in S. beali because of the large size of its host's seeds. We manipulated clutch size and examined its relationship to offspring fitness. Clutch size affected the survivorship of S. limbatus larvae(r ²=0.14), but had no detectable effect on the survivorship of S. beali larvae (r ²=0.04). Also, clutch size had a large effect on development time and body weight of S. limbatus, but not of S. beali. We discuss the implications of this result for the evolution of clutch size in S. limbatus and S. beali.     

THE EVOLUTIONARY GENETICS OF AN ADAPTIVE MATERNAL EFFECT: EGG SIZE PLASTICITY IN A SEED BEETLE

In many organisms, a female's environment provides a reliable indicator of the environmental conditions that her progeny will encounter. In such cases, maternal effects may evolve as mechanisms for transgenerational phenotypic plasticity whereby, in response to a predictive environmental cue, a mother can change the type of eggs that she makes or can program a developmental switch in her offspring, which produces offspring prepared for the environmental conditions predicted by the cue. One potentially common mechanism by which females manipulate the phenotype of their progeny is egg size plasticity, in which females vary egg size in response to environmental cues. We describe an experiment in which we quantify genetic variation in egg size and egg size plasticity in a seed beetle, Stator limbatus, and measure the genetic constraints on the evolution of egg size plasticity, quantified as the genetic correlation between the size of eggs laid across host plants. We found that genetic variation is present within populations for the size of eggs laid on seeds of two host plants (Acacia greggii and Cercidium floridum; h² ranged between 0.217 and 0.908), and that the heritability of egg size differed between populations and hosts (higher on A. greggii than on C. floridum). We also found that the evolution of egg size plasticity (the maternal effect) is in part constrained by a high genetic correlation across host plants (r_G > 0.6). However, the cross-environment genetic correlation is less than 1.0, which indicates that the size of eggs laid on these two hosts can diverge in response to natural selection and that egg size plasticity is thus capable of evolving in response to natural selection.     

Larviposition in the ovoviviparous blowfly Calliphora dubia

This study examined larviposition in Calliphora dubia Macquart (Diptera: Calliphoridae), an ovoviviparous blowfly of considerable forensic importance in Australia. Females in the field carried 22–83 live larvae, exhibiting a strong linear relationship between female size and the number of live larvae carried. Females took just over 1 min (mean 67.7 ± 7.7 s, n = 54) to larviposit live larvae on or near fresh liver in the laboratory. Females laid larvae at a mean rate of 1.2 ± 0.1 larvae/s, with the fastest rate being 3.4 larvae/s. Most females (70%) laid live larvae only, but 14% laid larvae and eggs at the same time and 16% laid eggs only (none of the eggs laid were viable). Females laying only live larvae laid a mean of 53.7 ± 2.3 larvae, whereas those laying only eggs laid a mean of 48.6 ± 2.8 eggs on each occasion. None of the eggs laid were viable. Most females (86%) laid all their larvae in a single spot, even if they engaged in several bouts of laying live larvae. Nearly one‐third of females did not lay all the live larvae in their ovisacs, but retained half of their complement of developed larvae. Females may be opting to spread their larvae across several carcasses in order to increase their survival and not to overcrowd small, ephemeral carcasses. The fact that a blowfly can lay either eggs or live larvae has enormous implications for the accurate determination of the post‐mortem interval (PMI) as the presence of larvae derived from eggs laid on the body add 6–18 h to the PMI. This paper represents the first report of the ability of female calliphorids to resorb some of their own live larvae.     

Population differences in host use by a seed-beetle: local adaptation, phenotypic plasticity and maternal effects

For insects that develop inside discrete hosts, both host size and host quality constrain offspring growth, influencing the evolution of body size and life history traits. Using a two-generation common garden experiment, we quantified the contribution of maternal and rearing hosts to differences in growth and life history traits between populations of the seed-feeding beetle Stator limbatus that use a large-seeded host, Acacia greggii, and a small-seeded host, Pseudosamanea guachapele. Populations differed genetically for all traits when beetles were raised in a common garden. Contrary to expectations from the local adaptation hypothesis, beetles from all populations were larger, developed faster and had higher survivorship when reared on seeds of A. greggii (the larger host), irrespective of their native host. We observed two host plant-mediated maternal effects: offspring matured sooner, regardless of their rearing host, when their mothers were reared on P. guachapele (this was not caused by an effect of rearing host on egg size), and females laid larger eggs on P. guachapele. This is the first study to document plasticity by S. limbatus in response to P. guachapele, suggesting that plasticity is an ancestral trait in S. limbatus that likely plays an important role in diet expansion. Although differences between populations in growth and life history traits are likely adaptations to their host plants, host-associated maternal effects, partly mediated by maternal egg size plasticity, influence growth and life history traits and likely play an important role in the evolution of the breadth of S. limbatus’ diet. More generally, phenotypic plasticity mediates the fitness consequences of using novel hosts, likely facilitating colonization of new hosts, but also buffering herbivores from selection post-colonization. Plasticity in response to novel versus normal hosts varied among our study populations such that disentangling the historical role of plasticity in mediating diet evolution requires the consideration of evolutionary history.     

Taking a chance on moths: oviposition by Delia flavifrons (Diptera: Anthomyiidae) on the flowers of bladder campion, Silene vulgaris (Caryophyllaceae)

Abstract.

• 1 Delia flavifrons Zetterstedt (Diptera: Anthomyiidae) visits flowers of Silene vulgaris (Moench) Garcke (Caryophyllaceae), where the adults feed, mate, lay their eggs, and the larvae feed on developing seeds. The objective of the study was to examine how an ovipositing female fly assures a food resource for her progeny.
• 2 Ovipositing females preferred young, non-pollinated flowers over older pollinated ones. The flies did not pollinate the flowers and survival of the larvae depended on the flowers being pollinated by moths.
• 3 Flowers containing fly eggs were pollinated more often than expected from chance, probably as a result of both flies and moths visiting particular flowers.
• 4 Eggs were laid singly, and multiple oviposition occurred randomly. Although most eggs hatched, only about half produced larvae that made their way into the fruits. As a result, the probability of competition arising from multiple oviposition may be reduced to such an extent that selection does not favour females that avoid flowers with conspecific eggs.
• 5 Moth larvae of the noctuid genus Hadena also feed on the seed pods of S. vulgaris and will kill any fly larvae they encounter. However, there was no deviation from random oviposition by the flies in relation to eggs laid by the moths, but the competitively weaker fly usually started to lay eggs towards the end of the moth's egg-laying period.
• 6 The relationship between Delia flavifrons and Silene vulgaris superficially parallels that for known pollinator/predator systems, but floral adaptations to hypothetically pollinating flies seems not to have taken place.

     

Tree colonization by the Asian longhorn beetle,Anoplophora glabripennis (Coleoptera: Cerambycidae): effect of habitat and tree suitability

Tree colonization and feeding activity of the invasive wood‐borer Asian longhorn beetle (Anoplophora glabripennis), an Asian pest introduced into North America and Europe, was studied in a newly invaded area in Italy. The hypothesis being tested was that the reproductive success of the insect depend on habitat type and tree suitability. Adult beetles were caged on branches of host and nonhost species, in both urban and forest habitats. Two months later, number and size of feeding patches on plant tissues, eggs laid, and surviving larvae were assessed. Bark concentration of C and N was also measured from the same trees. Results indicated that the mean area of plant tissues consumed by adult feeding was significantly larger on trees growing in forest than in urban habitat, although within the same habitat there were no differences between susceptible and nonsusceptible trees. ALB tree colonization, in terms of number of eggs laid and young larvae survival, was not affected by habitat while it was higher on susceptible trees. Although trees growing in forests had a lower nitrogen concentration, they allowed colonization rates similar to those of trees growing in the urban habitat. Hence, the amount of carbon and nitrogen did not fully explain tree suitability or habitat selection. We suggest compensatory feeding as a potential mechanism that might explain this peculiar situation, as supported by a more intensive feeding activity recorded on trees in the forest. Suitability of different trees may be due to other factors, such as secondary chemical compounds.     

Biology of Larinus curtus Hochhut (Coleoptera: Curculionidae), a European Weevil for Biological Control of Yellow Starthistle Centaurea solstitialis L (Asteraceae), in the United States

The biology of the weevil Larinus curtus Hochhut was studied in the field in northern Greece and in the laboratory in Rome, Italy, and in Thermi, near Thessaloniki, Greece. The species is univoltine, and adults overwinter in ground litter. Eggs are inserted into the flowers of yellow starthistle (Centaurea solstitialis L.) where the larvae feed mainly on developing achenes, destroying on average over 96% of the seeds in infested flowerheads. Overwintered adults lived up to 84 days, females laid up to 70 eggs each, eggs hatched 4.2 ± 0.6 days after being laid, larvae required 17 to 20 days to develop through the four instars, and pupal development required 4 to 5 days under laboratory conditions. Six percent of 360 seedheads collected on July 13 and 28, 1988 were infested with L. curtus larvae and up to 89% of the larvae were parasitized. The species is recommended for the biological control of C. solstitialis in the United States.     

Does insect mother know under what conditions it will make their offspring live?

According to the optimal oviposition theory, the larval success of insects depends on the oviposition site selection by females. Females are expected to choose a site with many resources and few competitors or predators to allow the best performance for their progeny, assuming that “mother knows best.” However, this is not systematically observed. The Aphidoletes aphidimyza larvae are generalist aphid predators and females consequently lay their eggs near or inside aphid colonies. The goal of this study was to investigate the impact of intraspecific competition on oviposition behavior of A. aphidimyza females. First, we counted the number of eggs laid by a female on a leaf with 20 aphids, in the presence of 0, 2, 4, or 6 conspecific eggs or in the absence of eggs but in presence of 3 virgin females. The same experiment was also performed under choice condition with 2 oviposition sites. Our results show that the presence of low densities of conspecific eggs, or the presence of conspecific females, have no significant impact on the number of eggs laid by A. aphidimyza females. One of the hypotheses advanced to explain these results is the advantages of conspecifics presence. At low densities, the presence of eggs on an oviposition site can indicate the suitability of the site for the females. The conspecific presence can also insure a dilution effect against predator and increase the presence of potential mating partners for this monogenic species.     

Reduced realised fecundity in the pine looper Bupalus piniarius caused by host plant defoliation

1. Whether the potential fecundity of herbivorous insects is realised or not may depend on female behaviour, which in turn may be influenced by host plant acceptability. Female Bupalus piniarius were observed to discriminate against needles growing out the year following defoliation (current‐year needles) of its host plant Pinus sylvestris. 2. It was hypothesised that the discriminatory behaviour was due to current‐year needles being less secure as a substrate. Field and laboratory experiments were designed to test this hypothesis and to estimate the discrepancy between potential and realised fecundity when females were offered defoliated branches. 3. In a laboratory oviposition experiment, B. piniarius females were exposed to branches bearing either current‐year needles only or both mature and current‐year needles. Daily oviposition rate, egg batch size, longevity, and mature eggs and fat retained at death were recorded for each female. In field experiments, the rate at which eggs dropped from expanding needles and the capacity of neonate larvae hatching from the dropped eggs to colonise a tree were assessed. 4. Significantly fewer eggs were laid when females were exposed to defoliated branches. 5. Twenty‐six and 16% of the eggs laid on current‐year needles dropped from the needles in 1998 and 1999 respectively, whereas no eggs dropped from mature needles in 1998 and only one egg (< 1%) dropped in 1999. 6. A very small proportion of larvae hatching on the forest floor (simulated egg drop) was able to recolonise host trees. 7. These results emphasise the importance of oviposition behaviour on realised fecundity when analysing insect population dynamics. In the case of B. piniarius, egg placement, although a minor detail during the normal course of events, became of key importance when defoliation deprived females of their preferred egg attachment site.     

Morphological features,development and reproduction of Melittobia acasta on Bombus terrestris

Morphological features, development and reproduction behavior of the parasite Melittobia acasta (Walker) were studied when reared on the pupae of the bumblebee Bombus terrestris L. in the laboratory under 23°C, 50% relative humidity and 12 h light : 12 h dark conditions. The parasites laid transparent white and elongated eggs. Newly hatched larval size and shape were very similar to eggs but they were identified by their body segments. Larvae increased their body size through moulting and transformed into a vermiform shape. Male pupae were shiny brown with dots. The female pupae were distinguished by their black shiny color, shorter size and the presence of compound eyes. Adult male pupae were dark brown and dwarf‐winged, whereas female pupae were macropterous and brachypterous. Reproduction took place by fertilization and also parthenogenetically. Mean fecundity within 5 days by mated (47.9 ± 30.5 female^?1) and virgin (7.4 ± 6.8 female^?1) females were statistically different. Mated females laid fertilized eggs that produced adult males or females, whereas virgin females laid unfertilized eggs that produced males. Development durations of the virgin female originated eggs, larvae, pupae and adults were statistically identical with those of mated females. The parasites were female‐biased and foundress number did not affect offspring sex ratio. This study shows that both mated and virgin females of M. acasta can produce many offspring on B. terrestris pupae within a short period, indicating that they are dangerous parasites of the bumblebee in a mass rearing system.     

Adaptive decision making or differential mortality: what causes offspring emergence in a gregarious parasitoid?

Hosts represent a limited resource for the developing offspring of parasitic insects laying eggs in or on spatially discrete resources like fruits, seeds, or other insects. The quality of hosts differs with respect to the value and amount of resources they provide for the feeding larvae. Accordingly, the size of a clutch of eggs laid on a given host should be a function of host quality, because severe competition between developing larvae can lead to increased mortality and/or decreased size of the offspring, both causing a fitness loss for the offspring and the mother. Therefore, females should be selected for the ability to estimate host quality and to adjust their clutch size accordingly. Using the parasitic wasp Nasonia vitripennis (Walker) (Hymenoptera: Pteromalidae) this study investigated the respective contribution of developmental mortality of offspring vs. the clutch size decision of the mother as a determinant of final offspring emergence per host. In addition, taking offspring size into account, the study examined the fitness consequences of female oviposition decisions. Developmental mortality was very low in all quality classes of hosts except previously frozen and thus dead host pupae. Females laid reduced clutch sizes on dead, previously parasitized, and smaller hosts. In contrast to offspring number, offspring size did not differ between host qualities. We conclude that females are able to sense the quality of a host and adjust the number of eggs they lay to mitigate larval competition.     

Protection of potato crop against Phthorimaea operculella (Zeller) infestation using frass extract of two noctuid insect pests under laboratory and storage simulation conditions

The oviposition deterrent effect of water extract of Spodoptera littoralis and Agrotis ipsilon larval frass on Phthorimaea operculella adult females was studied using two types of larval food “Natural host and Semi-artificial diet” under laboratory and storage simulation (semi-field) conditions. Extracted frass of fed larvae on semi-artificial diet showed complete oviposition deterrent effect at treatments with 4th, 5th and 6th instars of S. littoralis, also at treatments with 1st–3rd and 6th instars of A. ipsilon, while the same effect was observed when the larvae fed on castor oil leaves as a natural host only at treatment with frass extract of A. ipsilon 6th instar larvae. Presence of low amounts of phenols and flavonoids in water extract of A. ipsilon larval frass resulted in relatively more effect as oviposition deterrent to fertile adult females on treated oviposition sites, while the opposite effect was obtained in S. littoralis larval frass experiments. At semi-field experiments, the percentage reduction of laid eggs reached 100% after two?days at treatments with frass extracts of 4th and 5th S. littoralis larval instars and A. ipsilon 6th instar larvae fed on semi-artificial diet and/or castor oil leaves. Percentage reduction of laid eggs for untreated sacks reached 93.24 and 48.95% after 2 and 30?days, respectively, when placed between treated sacks, in comparison with the mean number of laid eggs for isolated control.     

Harem size and oviposition behaviour in a polygynous bark beetle

Abstract. 1. Harem polygyny can have fitness benefits and costs on females. In bark beetles of the genus Ips the latter may include within‐harem competition between larvae. However, earlier competition between females for male care and mating opportunities may also influence oviposition behaviour. There has been relatively little investigation into the relationship between harem size and initial egg output. The present study investigated this relationship in the bark beetle Ips grandicollis. 2. The measure of egg output used was the number of eggs in the gallery with the most eggs in each harem. Mean (±SE) harem size of 242 observed harems was 3.25 ± 0.10. A curvilinear relationship was found between egg output and harem size, with females in smaller harems (one to four females) laying more eggs with increased harem size. However, females in larger harems (five to seven females) laid fewer eggs as harem size increased. The optimal harem size (in terms of number of eggs laid) was close to four females. 3. We found no evidence from a behavioural assay that females could preferentially choose unmated males over mated males with harems of two females. Additionally, the distribution of harem sizes suggests that females distribute themselves among males randomly. 4. The results suggest that harem size has effects on female reproduction that extend beyond larval competition and influence patterns of oviposition. The mechanism that determines why egg laying is greatest at intermediate levels is unknown. There is no evidence that smaller harems belong to lower quality males, but females may adjust egg‐laying behaviour in large harems as a result of reduced male attendance or anticipated larval competition.     

Biology and life table studies of the oriental tobacco budworm,Helicoverpa assulta (Lepidoptera: Noctuidae), influenced by different larval diets

Development, survivorship, pupal weight, oviposition, and life table parameters of the oriental tobacco budworm, Helicoverpa assulta Guenée, were evaluated in the laboratory on an artificial diet, pepper (Capsicum frutescens L.), and tobacco (Nicotiana tobacum L.). We found that the average developmental time of immature stages was longest on tobacco (36.2 d), intermediate on pepper (34.4 d), and shortest on artificial diet (33.5 d). Immature survival from egg to pupa varied from 31% on tobacco, 43% on pepper, and 74% on artificial diet. Pupal weight ranged from 197.4 mg/pupa on tobacco, 233.1 mg/pupa on pepper and 253.4 mg/pupa on artificial diet. The average numbers of eggs laid by adults reared as larvae on the artificial diet, pepper, or tobacco were 614, 421 and 334 eggs/female, respectively. Numbers of remaining eggs in ovaries of the adult females reared as larvae on the artificial diet, pepper, or tobacco were 16, 26, and 42 eggs/female, respectively. The longevity of adult females developed from larvae reared on the three diets was not significantly different, whereas the longevity of male adults from the larvae reared on artificial diet was longer (16.8 d) than that for males reared on tobacco (13.8 d) and pepper (13.3 d). The intrinsic, finite, gross, and net rates of increase were highest for females reared as larvae on artificial diet, lowest for females emerging from larvae reared on tobacco, and intermediate for females emerging from larvae reared on pepper. Generation times and doubling time of H. assulta were shortest for larvae fed artificial diet, intermediate from larvae reared on pepper, and longest from larvae reared on tobacco. We concluded that the artificial diet was the most suitable larval diet of H. assulta followed by pepper, and tobacco.     

Egg size and composition in an ageing capital breeder – consequences for offspring performance

1. Egg size is often used as a proxy of egg quality although size and composition may vary, e.g. in insects egg size usually decreases as female ages. Whether this decrease in size reflects reduced concentrations of essential nutrients such as lipids and proteins of eggs laid by ageing females, or does reduced size per se explain often observed lower fitness of later laid eggs is poorly explored. 2. Egg properties were compared with fitness parameters of offspring laid on the first and fourth night during the oviposition period of a capital breeding moth, Cleorodes lichenaria (Hufnagel). The study aim was to explore whether decreased egg size is caused by decreased provisioning into later laid eggs measured as egg protein and lipid concentration, and whether it results in lower fitness of later laid offspring. 3. The fresh and dry weight of eggs decreased over the oviposition period, but the protein and lipid concentration remained constant. Survival of larvae was lower among the fourth night laid offspring on a low quality host Parmelia sulcata Taylor compared to a high quality host Ramalina fraxinea (L.) Ach. No differences were observed in egg fertility or hatchability, neonate survival without food and pupal mass between the offspring produced on different nights. 4. Decreased survival of offspring produced later was rather attributable to absolute provisioning (i.e. lower weight of eggs) than relative provisioning (i.e. decreased concentrations of nutrients in eggs). It is argued that lower survival of later laid smaller eggs on low quality diet is likely attributable to physical and chemical characteristics of host lichens and/or physical properties of tiny neonate larvae.     

Oviposition preference in the oligophagous tomato fruit fly, Neoceratitis cyanescens

We assessed the role of visual and olfactory cues on oviposition preference in the oligophagous tomato fruit fly, Neoceratitis cyanescens (Bezzi) (Diptera: Tephritidae). In a field survey, we evaluated the stage of susceptibility of field‐grown tomatoes by monitoring N. cyanescens infestations from fruit‐setting up to harvest, in relation to post‐flowering time, size, and visual properties of fruit. In two‐choice laboratory experiments, we tested the degree to which females use visual and olfactory cues to select their host plant for oviposition. In addition, we investigated the ability of flies to avoid fruit already infested by conspecific eggs or larvae, and the influence of natal host fruit on oviposition preference. Neoceratitis cyanescens females preferentially lay their eggs in small yellow‐green unripe fruit (2–3.5 cm diameter, 10–21 days post‐flowering). Damage to fruit was significantly affected by brightness and size properties. In laboratory experiments, females chose to lay their eggs in bright orange rather than yellow domes. On the sole basis of olfactory stimuli, females showed a significant preference for unripe vs. ripe host fruit, for unripe fruit vs. flowers or leaves, and for host vs. non‐host fruit (or control). However, colour interacted with odour as females dispatched their eggs equally between the yellow dome and the bright orange dome when unripe fruit of tomato was placed under the yellow dome vs. ripe fruit under the bright orange dome. When offered real ripe and unripe tomatoes, females preferred unripe tomatoes. Females significantly chose to lay eggs in non‐infested fruit when they were given the choice between these or fruit infested with larvae. In contrast, recent stings containing eggs did not deter females from laying eggs. Rather, they could have an attractive effect when deposited within <1 h. Regardless of their natal host plant, tomato or bugweed, N. cyanescens females laid significantly more eggs in a dome containing bugweed fruit. However, 15% of females originating from tomato laid eggs exclusively in the dome with tomato, against 3% of females originating from bugweed.     

The biology of the ground beetle Harpalus rufipes in a strawberry field in Northumberland

Harpalus rufipes (Degeer) was studied in a strawberry plot in Northumberland from 1973 to 1978 by pitfall trapping, and in the laboratory. Adults were active from April until November. Overwintered male beetles predominated at the beginning of each season until May, followed by overwintered females in June and July. Newly emerged, mainly female, beetles were active from August onwards. Overwintered females matured during early summer and laid eggs in August with a fecundity of 10–15 eggs/female. In the laboratory about 30% of beetles survived from one breeding season to the next. First- and second-instar larvae were caught in pitfall traps in autumn; in the laboratory they made approximately vertical burrows in which they stored seeds taken from the soil surface. Third-instar larvae fed on these seeds and were not active on the surface. Preferred seeds were those of grasses and Chenopodium album L. Larvae were usually aggregated in the soil at densities of 3–20/m².