Fecundity and egg viability in relation to female body size in Neodiprion sertifer (Hymenoptera: Diprionidae)

Relationships between female size and fecundity, egg viability and embryonal development were studied in Neoriprion sertifer. Sawfly females, reared from the 1st instar larvae on Scots pine needles heavily affected by industrial pollutants or on relatively unpolluted needles, were induced to lay eggs parthenogenetically in the field in Finland. Females reared on polluted needles were smaller and oviposited an average of 68.8 eggs, and those reared on unpolluted needles 79.4 eggs per female. However, the higher egg viability in the former group masked the effect of reduced female size. This compensation was primarily related to the size of the females, larger individuals producing relatively fewer viable eggs in both groups.     

Reduction of ethylene emission from Scots pine elicited by insect egg secretion

Pinus sylvestris L. is known to activate indirect defence in response to attack by an herbivorous sawfly. Egg deposition by the sawfly Diprion pini L. induces pine to release, three days after egg laying, locally and systemically terpenoid volatiles that attract parasitoids to kill the eggs. The elicitor of the pine's response is located in the sawfly's oviduct secretion enveloping the eggs after deposition. Application of this secretion on twigs with artificially conducted ovipositional woundings mimics the effects of egg deposition. Furthermore, jasmonic acid (JA) induces a volatile pattern similar, but not identical, to the one induced by egg deposition. To gain deeper insight into the transduction of plant signals induced by herbivore egg deposition, it was investigated whether ethylene emission from pine is affected by sawfly egg deposition. Systemically induced ethylene emission from differently treated pine twigs was monitored for a period of 3 d after treatment. Ethylene emissions from untreated control twigs were compared with those from twigs treated as follows: (i) sawfly egg secretion [=oviduct secretion (OVI)] was transferred on artificially wounded pine needles (attractive volatiles), (ii) needles were artificially wounded (non-attractive volatiles), and (iii) the twig was supplied with JA (attractive volatiles). Ethylene emission from systemically OVI-induced twigs was significantly lower than from untreated controls, whereas artificial wounding had no detectable effect. JA-treated twigs released much more ethylene and showed higher variability of ethylene emission than artificially wounded twigs and OVI-treated ones. Ethylene emissions from pine after the various treatments studied here are discussed with respect to known effects of insect feeding on ethylene release from plants.     

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.     

Significant mortality of eggs and young larvae of two pine processionary moth species due to the entomopathogenic fungus Metarhizium brunneum

Bioassays were conducted to determine the susceptibility of egg masses and young larvae of two pine processionary moth species, Thaumetopoea pityocampa and Thaumetopoea wilkinsoni, to two strains (ARSEF4556, V275) of the entomopathogenic fungus Metarhizium brunneum. Mortality of treated eggs by both strains ranged from 96% to 99% but not all of this was caused by M. brunneum since control groups also experienced egg mortality due to saprophytic fungi. Still, larvae hatched in the laboratory from eggs treated with M. brunneum were all killed by this fungus, acquiring M. brunneum conidia, whereas larval mortality was 0% in the control groups. Young larvae of both pine processionary moth species were also highly susceptible to ARSEF4556 and V275 with larval mortality ranging between 94% and 100%, 8 days post-inoculation, with the vast majority of larvae being killed within the first 2–4 days. Larval mortality was dose dependent. Results were consistent across the two pine processionary moth species, showing that the pathogenicity of M. brunneum to both eggs and young larvae might be promising for biological control of these insect pests. The study also showed that non-target parasitoids of pine processionary moth eggs were also susceptible to M. brunneum. Further work is required to understand and reduce the M. brunneum effect on non-target insects.     

Effect of forest stand type on host plant quality and direct and indirect effects on pine sawfly performance

1. Observed lower levels of herbivory in mixed compared with monoculture stands have been hypothesized to depend on top-down forces, through higher predation pressure by natural enemies or through bottom-up mechanisms through plant quality effects on herbivore performance.
2. In this study, we compared the performance measured as host plant induced mortality, cocoon weight, and predation mortality of the European pine sawfly Neodiprion sertifer (Geoffroy) (Hymenoptera, Diprionidae) in mixed and monoculture forest stands.
3. We did not observe a difference in host plant induced mortality, cocoon weight, or predation mortality between mixed and monoculture forest stands. We did find an effect of local conditions around each experimental tree on pine sawfly performance. For example, the nitrogen content of pine needles is negatively affected by the proportion of pine around the experimental tree, which in turn increases the survival of sawfly larvae.
4. The results suggest that local conditions around individual trees are more important for the performance of the European pine sawfly than stand type, i.e. mixed or monoculture plant stands.
5. We conclude that the ongoing trend for diversification within commercial forestry calls for more research where the effects of both bottom-up and top-down effects are studied at several spatial scales.

     

The role of low-level ozone exposure and mycorrhizas in chemical quality and insect herbivore performance on Scots pine seedlings

The effects of low‐level ozone exposure and suppression of natural mycorrhizas on the above‐ground chemical quality of Scots pine (Pinus sylvestris L.) needles and insect herbivore performance were studied in a two‐year field experiment. Seedlings were fumigated with the ozone doses 1.5–1.7 times the ambient, and natural mycorrhizal infection level was about 35% reduced in roots with fungicide propiconazole. On ozone‐exposed seedlings the mean relative growth rate (MRGR) of Lygus rugulipennis Popp. nymphs was lower than on ambient ozone seedlings, but Gilpinia pallida Klug sawfly larvae grew better on elevated ozone seedlings than on ambient ozone seedlings. MRGR of Schizolachnus pineti Fabr. and Cinara pinea L. aphid nymphs or Neodiprion sertifer Geoffr. sawfly larvae or the oviposition of L. rugulipennis and N. sertifer were not affected by ozone exposure. Although ozone exposure did not affect total phenolics, total terpene, total or individual resin acid, total free amino acid, nutrient or sugar concentrations in needles, MRGR of L. rugulipennis positively correlated with total terpenes and MRGR of G. pallida positively with total amino acids. In addition, ozone exposure increased serine and proline concentration and marginally also starch concentration in needles. When mycorrhizas were reduced with fungicide, only MRGR of L. rugulipennis nymphs increased, but performance of other insect herbivores studied was not changed. However, number of L. rugulipennis eggs correlated positively with mycorrhizal infection level and also with total sugars. Reduction of mycorrhizas did not strongly affect the concentrations of analysed compounds in needles, because only phosphorus and potassium and some individual resin acids were reduced by fungicide treatment. These results suggest that low‐level ozone exposure and moderately declined mycorrhizal infection do not drastically affect either the above‐ground chemical quality of Scots pine seedlings or performance of studied insect herbivores.     

Effects of group size and pine defence chemicals on Diprionid sawfly survival against ant predation

The defence chemicals and behavioural adaptations (gregariousness and active defensive behaviour) of pine sawfly larvae may be effective against ant predation. However, previous studies have tested their defences against very few species of ants, and few experiments have explored ant predation in nature. We studied how larval group size (groups of 5 and 20 in Neodiprion sertifer and 10, 20 and 40 in Diprion pini) and variation in levels of defence chemicals in the host tree (Scots pine, Pinus sylvestris) affect the survival of sawfly larvae. Food preference experiments showed that ants do eat sawfly larvae, although they are not their most preferred food item. According to our results, ant predation significantly increases the mortality rate of sawfly larvae. Larval mortality was minor on pine tree branches where ant traffic was excluded. We also found that a high resin acid concentration in the host tree significantly decreased the mortality of D. pini larvae when ants were present. However, there was no such relationship between the chemical concentrations of the host tree and larval mortality for N. sertifer. Surprisingly, grouping did not help sawfly larvae against ant predation. Mortality risk was the same for all group sizes. The results of the study seemingly contradict previous understanding of the effectiveness of defence mechanisms of pine sawfly against ant predation, and suggest that ants (Formica exsecta in particular) are effective predators of sawfly larvae.An erratum to this article can be found at     

Hatching and viability of rotifer diapausing eggs collected from pond sediments

1. Planktonic rotifers inhabiting variable environments produce diapausing eggs that accumulate in the sediment of lakes and ponds, forming egg banks that may withstand adverse periods. A common assumption in zooplankton diapausing egg bank studies is to count as viable all eggs in the sediment that look healthy. This assumption should be challenged by asking how effectively ‘healthy‐looking’ eggs represent viable eggs. 2. In this study, viability of more than 1100 ‘healthy‐looking’ diapausing eggs belonging to the Brachionus plicatilis species complex was assessed in a laboratory hatching experiment. Eggs were collected at different depths from sediment cores obtained from 15 ponds located in coastal and inland areas of Eastern Spain. 3. Only approximately one half of the ‘healthy‐looking’ diapausing eggs hatched after incubation in experimental conditions. Almost all the hatchlings (99.4%) survived to maturity. The proportion of ‘healthy‐looking’ diapausing eggs that hatched varied among areas and among ponds within area, and substantially declined with sediment depth. Most of the hatchlings (88%) were obtained from the uppermost 2 cm of sediment. ‘Healthy‐looking’ eggs from upper sediment layers hatched after significantly shorter incubation times than eggs recovered from deeper layers. 4. Both decreased hatching success and increased incubation time for hatching with sediment depth suggest that older ‘healthy‐looking’ eggs are less responsive to hatching stimuli and could become unviable. However, the strong correlation found between the number of ‘healthy‐looking’ eggs and the number of hatchlings indicates that the abundance of ‘healthy‐looking’ eggs is a good index of egg bank viability.     

To hatch or not to hatch? Egg hatch response to larval density and to larval contact in a treehole mosquito

Abstract.

• 1 We investigated the effect on egg hatch of exposure to: (1) varying larval density, and (2) larval contact in Aedes triseriatus Say (Diptera: Culicidae). For 2 days in the laboratory we submerged eggs into a treehole water medium containing 0 (control), 4, 12 or 24 larvae that could either contact the eggs directly or were separated from them by a screen. Following treatment, abundance of microorganisms on the egg surfaces, a food source for newly hatched larvae and a proposed hatching stimulus, was assessed by counts made from serial dilutions of samples.
• 2 We discovered a complex hatching response to larval contact and to larval density, and an interaction between these two factors in their effect on microbial growth. Hatching was inhibited in the 0-larva control, even though microorganisms grew abundantly on the eggs. Hatch rate, as well as microbial counts, were high for eggs in direct contact with 4 larvae. As density increased in the larval contact treatment, microorganisms disappeared from the egg surfaces and hatch rate declined.
• 3 When protected from larval grazing, eggs supported numerous microbial colonies irrespective of larval density. In contrast to the contact treatment group, egg hatch increased with increasing larval density. These observations suggest that the combination of microbial growth and a larval factor stimulates hatch. This hatching response may have evolved because both abundant microorganisms and numerous larvae reflect a habitat of good quality.

     

Simulated acid rain reduces the susceptibility of the European pine sawfly (Neodiprion sertifer) to its nuclear polyhedrosis virus

Summary The study dealt with the effect of simulated acid rain (both H₂SO₄ and HNO₃; acidities of pH 4 and pH 3) on the susceptibility of the larvae of Neodiprion sertifer to its nuclear polyhedrosis virus. Scots pines growing in a subarctic area with low ambient pollution levels were irrigated with simulated acid rain during two summers. Neodiprion larvae fed with foliage from the experimental trees were infected with a dilute virus suspension. The acid treatment of host trees had a significant effect on the proportion of virus-treated larvae alive 16 days after the virus application: there were almost no differences between the controls and the pH 4 irrigation group, but on the needles of pH 3-treated trees larval survival was twice as high as with other treatments. The direct spraying of acid water on the needles before they were fed to the larvae did not significantly affect the survival of virus infected larvae. Our results suggest that acid rain may reduce the susceptibility of Neodiprion larvae to virus disease via changes in the quality of pine foliage.     

Impact of the parasitoid Lathrolestes ensator (Hym., Ichneumonidae, Ctenopelmatinae) as antagonist of apple sawfly Hoplocampa testudinea (Hym., Tenthredinidae)

Abstract: Lathrolestes ensator (Brauns) was the only parasitoid species reared from larvae of apple sawfly Hoplocampa testudinea (Klug) collected in Dutch orchards. The life history parameters of apple sawfly and its parasitoid L. ensator were investigated in field and semi‐field experiments. Usually, the adult parasitoids emerge in synchrony with the preferred host stage, the second instar sawfly larvae. Parasitoid females carried 120–175 eggs, but never laid more than half this number. The mortality of sawfly eggs and larvae varied from 75 and 99%. The eggs and mining larvae are most vulnerable, as the older larvae survive for 90% on average. Three factors at least determine how many of the descending larvae become adult sawfly or parasitoid next spring. The failure of descending larvae to construct a cocoon varied from 7 to 31% and was highest in clay soil. Probably a similar mechanical hindrance prohibited more emerging adults, of both sawfly and L. ensator, from attaining the surface in heavier soils next spring. Relatively more sawfly prepupae than parasitoids died in the cocoon stage, from fungus disease or other causes, but more parasitoids than hosts stayed in prolonged diapause. Mainly due to this last factor, the overall result was a decrease of the parasitoid : host ratio during the first season underground. A life table based on survival rates during various life stages yields a net reproductive rate of up to 2.4 daughters per female sawfly. It indicates that 60% of the sawfly larvae need to be parasitized to stop population growth, or correspondingly less when more than 75% of the host eggs and young larvae are killed by predators or other causes.     

Reproductive bionomics of the soft tick, Ornithodoros turicata (Acari: Argasidae)

The effects of different temperatures and relative humidities (RHs) were tested on various reproductive parameters of Ornithodoros turicata, an argasid tick that inhabits gopher tortoise burrows in Florida, USA. The pre-oviposition, oviposition and incubation periods of the ticks decreased as temperature increased. These periods were also affected by the RH. The number of eggs oviposited was affected significantly by the combined effect of temperature and RH. Fewer eggs were laid by ticks in the 24°C regimes and the 27°C/95%RH regime compared to those in the other temperature/RH groups. There was an inverse relationship between the number of eggs oviposited and the percentage of hatched larvae that was correlated with the temperature and RH. Ticks reared at 27°C/90%RH and 30°C/90%RH laid more eggs than those reared in the other combinations of temperature and humidity but fewer larvae hatched from these eggs. The reproductive fitness index (RFI) values were highest in females held in the 24°C groups and the 30°C/95%RH group, although significantly more larvae hatched at the lower temperatures. The optimum reproductive conditions for O. turicata under laboratory conditions appear to be 24°C and 90–95%RH. While mating occurred at all temperatures, none of the females laid eggs at 22°C. The ticks may move preferentially to low temperatures when not feeding to remain above the critical equilibrium humidity and/or below the critical metabolic level necessary for prolonged survival. However, most female ticks oviposited after 45 days when moved to 27°C/95%RH. Ornithodoros turicata females may have a limited capability to delay oviposition until an optimal microenvironment for egg deposition can be located in the burrow.     

A case of abnormal size in eggs and larvae of the Taiga tick,Ixodes persulcatus (Acari: Ixodidae)

Egg batches laid by females of Ixodes persulcatus (maintained under a photoperiod of L:D 20:4 before feeding) contained eggs of two sizes: numerous normal eggs of 0.578 × 0.421 mm and some giant eggs of 0.776×0.515 mm. Giant eggs were 32–34% greater in length and 22–24% greater in width than normal eggs. Females maintained under a short-day photoperiod (L:D 12:12) laid only normal eggs (0.588×0.417 mm). There were no differences between these eggs in viability and developmental time, but larvae hatched from giant eggs were larger (length of body 23–28% greater and of gnathosoma 12–15% greater) than larvae from normal eggs. This dimorphism of eggs and larvae was not sex determined, but may be a maternal photoperiodic effect.     

Simultaneous maturation of two discrete oocyte batches in ayu

Ayu Plecoglossus altivelis including amphidromous and landlocked populations, are known for batch spawning. Unusual clutches at ovulation were observed from rearing stocks that were naturally induced to sexual maturation. Two peaks occurred simultaneously in the frequency distribution of egg size, compared to normal fish, which have clutches with unimodal size distribution. The eggs in the small size class could be fertilized, and small-sized larvae hatched from these eggs. To explain the alteration in egg size through divergence from amphidromous to landlocked populations, the role of batch allocation among oocytes is discussed.     

Egg cannibalism in Helicoverpa armigera on sorghum and pigeonpea

Egg cannibalism by Helicoverpa armigeraHübner (Lepidoptera: Noctuidae) larvae was studied in the laboratory and in the field. In laboratory experiments, first instars were exposed to increasing densities of H. armigera eggs on sorghum, Sorghum bicolor (L.) Moench, and pigeonpea, Cajanus cajan (L.) Millsp. The number of eggs eaten per larva increasedsignificantly as egg availability increased on both sorghum and pigeonpea. In small cages 21–37% of eggs were eaten on sorghum, 4–12%on pigeonpea. Plant feeding declinedsignificantly on both sorghum and pigeonpea asegg density increased. Cannibalism was greateron sorghum than on pigeonpea while plantfeeding was greater on pigeonpea than onsorghum. Only around 8% of eggs were eaten inlarger cages with sorghum. The response toincreasing egg availability in all experimentswas linear. Immunoassay with an anti-vitellinmonoclonal antibody showed that egg cannibalismoccurs on pigeonpea under field conditions.Seven percent of all larvae had egg protein intheir gut. Cannibalism may make a significantcontribution to H. armigera populationsuppression.     

Field Assessment of Adhesion and Hatch ofChrysoperlaEggs Mechanically Applied in Liquid Carriers

A mechanical technique was evaluated for releasing green lacewing eggs in liquid suspensions. Deposited eggs were enclosed within a circle of nonpoisonous adhesive to protect them from predation and to prevent escape of hatched larvae. Released eggs were monitored daily for 5 days after release by measuring three response variables: adhesion rate of eggs to foliage, hatch rate of eggs, and “yield” of larvae from discharged eggs; “yield” was the product of egg adhesion and egg hatch. Factors tested were: egg conditioning prior to release (incubated or refrigerated), carrier (distilled water or commercial carrier solution), application technique (mechanical or hand application), and row facing (North or South). Release technique did not significantly effect egg hatch on any day. Conditioning eggs prior to release had the greatest effect on hatch of eggs and resulting yield of larvae during the 5-day monitoring period. Carrier had a significant effect on adhesion of eggs to leaves and hatch of eggs. Commercial carrier solution increased egg adhesion but decreased egg hatch compared to water. Overall mean yield of larvae from incubated eggs distributed mechanically was not significantly different for eggs suspended in water (36.4% on day 5 post-release) and for eggs suspended in commercial carrier solution (36.1% on day 5 post-release). Hand-applied eggs had a higher hatch and subsequent yield of larvae than mechanically released eggs; however, the hand technique was labor intensive.     

Long-term effects of exogenous methyl jasmonate application on Scots pine (Pinus sylvestris) needle chemical defence and diprionid sawfly performance

Scots pine [ Pinus sylvestris L. (Pinaceae)] trees with four different seed origins were exposed to exogenous applications of the elicitor, methyl jasmonate (MeJA), for three consecutive years. We studied the effects of MeJA on needle chemistry (including monoterpenes, sesquiterpenes, and tricyclic resin acids), plant growth, and the performance of two diprionid sawflies, the European pine sawfly ( Neodiprion sertifer Geoffr.) and the common pine sawfly ( Diprion pini L.) (both Hymenoptera: Diprionidae). In general, foliar MeJA application affected the whole range of needle secondary chemistry with significantly higher concentrations of two monoterpenes, β-pinene and limonene, in particular. Furthermore, for some seed origins the growth rates of N. sertifer and D. pini larvae were lower on needles of MeJA-treated plants with either high total terpene or high resin acid concentrations. However, inconsistencies in diprionid sawfly performance within each studied Scots pine origin suggest genetic variance in needle secondary chemistry. The differences between selected seed origins and notably variable responses to MeJA application imply that adaptation of the seed to new conditions may have had an impact on secondary chemistry and, thus, on insect performance. Finally, our results suggest that modification of Scots pine defence by a low-concentration exogenous elicitor affects the production of terpenoids in the newly growing needles, leading to poorer pine sawfly performance in origins with high terpenoid content, while not harming the growth of Scots pine trees.     

Differentiated egg size of the cannibalistic salamander <Emphasis Type="Italic">Hynobius retardatus</Emphasis>

Larvae of the salamander, Hynobius retardatus, are carnivorous, and even though there are two morphs, a typical morph and a broad-headed or “cannibal” morph, both are cannibalistic. They also sometimes eat other large prey, for example larvae of the frog, Rana pirica. In natural habitats, use of both conspecific and R. pirica larvae as food may contribute more strongly to high survival and substantially to fitness when larval densities are higher, because early-stage H. retardatus larvae sometimes experience scarcity of their typical prey. In cannibalistic oviparous amphibians, larger individuals that developed from larger eggs can more efficiently catch and consume larger prey and thus their survival may be better than that of smaller individuals developed from smaller eggs. Populations might therefore diverge in respect of egg size in response to variation in the density of conspecific and R. pirica larvae in natural ponds, with eggs being larger when larval density is higher. I examined how variance in hatchling size correlated with the incidence of cannibalism, and whether increasing larval density in natural ponds correlated with increasing egg size. Variance in initial larval body size facilitated cannibalism, and egg size increased as larval density in the ponds increased. In ponds with high larval density, where cannibalism and large prey consumption is a critical factor in offspring fitness, the production of fewer clutches with larger eggs, and thus of fewer and larger offspring, results in greater maternal fitness. Variation among the mean egg size in populations is likely to represent a shift in optimum egg size across larval density gradients.