Leaf age and larval performance of the leaf beetle Paropsis atomaria

ABSTRACT.

• 1 Larval performance of the leaf beetle Paropsis atomaria Oliver was determined for larvae raised on both new and mature leaves of Eucalyptus blakelyi Maiden. Larvae were transferred to mature leaves at different ages; control larvae stayed on new leaves through all instars.
• 2 Only larvae reared on new leaves through the third instar survived to pupate on mature leaves; developmental time was prolonged by 20% and pupal weight was reduced by 50% in these larvae compared with larvae reared entirely on new leaves. Almost all larvae died when transferred to mature leaves as first, second or third instars.
• 3 Low survival and slow development on mature leaves was mainly due to failure by larvae to feed. Larvae palpated leaves and could discriminate among leaf ages immediately, without biting into the leaf tissue.
• 4 New leaves had higher concentrations of oil and tannins than old leaves, while there were no significant differences in nitrogen concentrations in the two types of leaves. Mature leaves were more than 3 times tougher than new leaves, in terms of g mm^?2 of penetrometer force.
• 5 In drought years E. blakelyi may not produce sufficient new leaves to supply specialist herbivores with their preferred food resource. We infer that drought years reduce P. atomaria larval performance significantly, and influence the population dynamics of the insect.

     

Development of the Periphytic Community on Artificial Substrates in Fish Ponds

The development of the periphytic algae and macrofauna was examined in two fish ponds. 43 genera of algae (mainly diatoms) were found on the substrates, and the maximum of algae was reached on the 10th day. The periphytic macrofauna consisted of 37 taxonomical groups (mainly Chironomidae). Larvae of Cricotopus intersectus were the most abundant species. Since the 2nd day of exposure, the number of Cricotopus larvae strongly increased by immigration of 2nd instars until the maximum was reached on the 15th day. No further significant oscillations were found in either the algal or animal component until the 75th day of exposure. The field observations and laboratory experiments confirmed that:

• (1) The distribution of frequencies of Cricotopus larvae found on 200 cm² surfaces could be approximated by the normal distribution.
• (2) The density of the algal periphyton is controlled by abiotic factors, not by grazing by chironomid larvae.
• (3) The Cricotopus larvae of the 2nd instar colonize the substrate only after the algal film has been formed. Later instars are sedentary and do not colonize new substrate (except under stress conditions).
• (4) Food and predation do not seem to be the factors controlling the Cricotopus population density. The main factor is considered to be the territorial behaviour of the larvae.

     

Factors affecting litter processing by Anisocentropus kirramus (Trichoptera: Calamoceratidae) from an Australian tropical rainforest stream

• 1 Larvae of the caddisfly Anisocentropus kirramus are common leaf shredders in rainforest streams in tropical Queensland. Laboratory experiments were undertaken to examine the effects of (a) differences in leaf species, age and degree of conditioning, (b) leaf preferences, (c) temperature and (d) larval stage on processing of leaf litter by A. kirramus.
• 2 Leaf species (five tested), age (fresh v senescent) and condition (conditioned in stream v unconditioned) each had a significant effect on the amount of leaf material that was processed. For most species, conditioned leaves were processed faster but for one species unconditioned leaves were processed faster. Senescent leaves were processed more rapidly than green leaves in three species but not in the other two. Given a choice of leaf types A. kirramus actively selected leaves that were processed faster; no preference was shown between two different ‘fast’ leaves.
• 3 Processing occurred at all temperatures tested (10, 18 and 25°C), with the processing rate increasing with temperature. Younger instar larvae processed leaves at a greater rate per unit body weight (up to 343% day^?1) than older instars. Final instar (5) larvae were capable of processing some tough leaves that younger instars did not process.
• 4 In rainforest streams, processing of leaves by A. kirramus takes place throughout the year. Its ability to process green leaves is important because of the high input of fresh green leaves into tropical streams, and because of the severe depletion of the supply of conditioned leaves and fine detritus after floods.

     

Larval aestivation and direct development as alternative strategies in the speckled wood butterfly, Pararge aegeria, in Sweden

ABSTRACT.

• 1 Sweden has two disjunct populations of the speckled wood butterfly, Pararge aegeria L. The southern population has two generations per year but the central Swedish population is univoltine. When rearing larvae from central Sweden under normal photoperiodic conditions but at temperatures slightly above the ambient, 42% of the larvae developed directly and produced a second generation of adults the same summer. The egg—larval development time of the directly developing individuals was about 40 days, whereas that of the individuals developing along the univoltine pathway was about 100 days.
• 2 Larvae of the central Swedish population normally aestivate during part of the summer even though abundant food is available. In the closely related Lasiommata petropolitana F., which is the only Swedish satyrid that overwinters in the pupal stage besides P.aegeria, larvae do not aestivate, indicating that there does not seem to be any obligatory association between pupal hibernation and larval aestivation.
• 3 Development rates of aestivating and directly developing P.aegeria are equal up to the third larval instar. During the third and fourth instars, however, the development rate of aestivating individuals is retarded and females also have an additional fifth instar.
• 4 Since the central Swedish P.aegeria have the capacity to develop directly, and the southern Swedish ones have the capacity to aestivate, the evidence indicates that the outcome of the cost/benefit balance of univoltine versus bivoltine development differs between the two areas.

     

Studies on Chironomidae in experimental recirculating stream systems. II. The growth, development and production of a spring generation of Orthocladius (Euorthodadius) calvus Pinder

SUMMARY.

• 1 Orthodadius (Euorthodadius) calvus Pinder. similar to Orthocladius (Euorthodadius) thienemanni Kieffer. colonized a new gravel substratum in two recirculating stream channels. A maximum population density of 68.621 m^-2 was attained after only 16 days. This had fallen to a negligible density by the thirty-third day.
• 2 Some recruitment occurred over most of the study period (April-May 1981) and no single sharply defined cohort was evident. There was a large range of body lengths within each instar. and considerable overlap between instars. The population density estimates for instars I and II were low compared with instars III and IV.
• 3 The relationship between instantaneous growth rate (g) and geometric mean length indicated that growth was best described by a Gompertz curve. Growth rate decreased with increasing length from a value of about 40% length day^-1 at 2 mm body length to about 5% length day^-1 at 9 mm.
• 4 Growth rates for individual larvae, kept in culture, were very variable with maximum rates close to the values determined from the field data. Mean duration of larval life was 16 days.
• 5 Estimates of production for the study period ranged from 13.5 g dry wt m^-2 (Channel III, size-frequency method) to 34.2 g dry wt m^-2 (Channel II, Allen's graphical method, values corrected for non-linear growth).
• 6 Gut contents were estimated to represent about 55% of the total weight therefore production values should be reduced by this amount.

     

Assimilation of carbon and nitrogen from pollen and nectar by a predaceous larva and its effects on growth and development

Abstract. 1. Predaceous insects may benefit from feeding on non‐prey foods, such as pollen, nectar, and honeydew, because they can provide nutrients that help maintain metabolism and enhance overall nutrient intake. Yet, the extent to which predaceous insects can assimilate non‐prey food and the importance of diet mixing during particular life history stages is poorly understood. In this study the relative contribution of an omnivorous diet to the growth and survivorship of a predaceous larva was tested in a hypothetical situation in which nutritionally optimal prey was not available. The study system comprised a predaceous larva (second‐ and third‐instar larvae of the green lacewing Chrysoperla carnea), nutritionally poor prey (larvae of Drosophila melanogaster), and non‐prey food (pollen suspension, a mixture of bee pollen and artificial nectar (1 M sucrose solution)). Chrysoperla carnea larvae in the mixed diet treatment were provided with both Drosophila larvae and pollen suspension, while those reared on the prey and non‐prey diet treatments received only Drosophila larvae or pollen suspension respectively. 2. The inclusion of pollen and sucrose in their diet enhanced the growth of C. carnea larvae. Second instars reared on the mixed diet developed significantly faster than their cohorts reared on the prey diet, however third instars reared on the mixed diet did not develop faster than their cohorts reared on the prey diet. Larvae reared on the mixed diet became larger adults than did those reared on either the prey or non‐prey diets. Third instars reared on the non‐prey diet completed their development while second instars in the non‐prey diet treatment failed to pupate. 3. Stable isotope analysis indicated that the larvae obtained most of their carbon (55–73%) and nitrogen (71–73%) from Drosophila but acquired only a minor amount of carbon (2–5%) and nitrogen (3–11%) from pollen. Larvae reared on the mixed and non‐prey diets acquired a relatively significant amount of carbon (23–51%) from sucrose. 4. A model, which included a novel fractionation factor to account for the isotopic effect of metamorphosis, was developed to explain the proportion of larval growth attributable to each diet item. It explained the adult δ¹³C values to within 0.2‰ and adult δ¹⁵N values to within 0.7‰ in all treatments. 5. Adults fed ¹⁵N‐labelled pollen as larvae retained the ¹⁵N signal of the pollen as adults. 6. The collective results of this study support the view that, despite their dependence on prey arthropods to obtain most of their dietary nitrogen, omnivorous lacewing larvae can enhance their growth and development by supplementing their diets with alternative non‐prey food resources. This finding is consistent with the notion that omnivory has evolved as a feeding strategy to acquire both additional nitrogen as well as trace nutrients.     

The Association of the Stream Caddisfly Lasiocephala basalis (Kol.) (Trichoptera: Lepidostomatidae) with Wood

A population of the lepidostomatid caddisfly, Lasiocephala basalis, was studied in a third order stream, the Ohe, in Hesse, Germany. L. basalis showed an intimate association with wood debris in all of its aquatic life stages. Larvae exhibited a significant preference for organic substrates such as woody debris, leaves, and submerged alder roots. Gut content analysis revealed leaf material and wood fragments as the dominant larval food items classifying larvae as facultative xylophagous. Larval dominance on wood during spring corresponded to minimal leaf litter abundance. In winter, during periods of high discharge, the most important larval habitat was the submerged alder roots at the stream margin, thus acting as flow refugia. Most pupal cases were attached to submerged logs and roots. In addition, most of the egg masses were found on these two substrates. The role of L. basalis larvae in wood degradation is discussed.     

The drag coefficient of cased caddis larvae from running waters: experimental determination and ecological applications

• 1 Drag coefficients of cylindrical-case caddis larvae from running waters were investigated in the laboratory using an artificial stream channel.
• 2 Dead larvae of the limnephilids Allogamus auricollis (first to fifth instars), fifth instars of Potamophylax cingulatus, Chaetapteryx fusca, Drusus monticola, Metanoea rhaetica and fourth instars of the brachycentrid Micrasema minimum, were exposed to different current speeds. When the heads of the larvae were directed towards the water flow (frontal position), the current necessary to dislodge the larvae ranged from 3.00cms-¹ (A. auricollis, first instar) to 70.50cms-¹ (P. cingulatus). With flow normal to the long axis of the case (lateral position), these speeds ranged from 2.20cms-¹ (M. minimum) to 20.80cms-¹ (P. cingulatus).
• 3 In frontal position, individual Reynolds numbers at the moment of dislodgement ranged from 74 (A. auricollis, first instars) to 14100 (P. cingulatus), and from 14 (M. minimum) to 1143 (P. cingulatus) in lateral position. Regression equations correlating case length, mean case width or fresh weight with Reynolds numbers at the moment of dislodgement (frontal and lateral position) were very highly significant with r²0.91.
• 4 For the range of Reynolds numbers given above, the drag coefficient varied between 5.05 (A. auricollis, first instars) and 0.26 (P. cingulatus) in frontal case position and from 2.97 (M. minimum) to 0.69 (C. fusca) in lateral case position. Furthermore, the relationship between Reynolds number and drag coefficient was found to be linear on a In/In scale for both frontal and lateral case position.

     

The food and daily food consumption of lacustrine 0+ perch, Perca fluviatilis L.

SUMMARY.

• 1 The diet and daily food consumption (C_t) of 0+ perch were investigated in two shallow Scottish lakes.
• 2 The first food comprised the rotifer Keratella sp., Volvox sp., copepod nauplii and stage I copepodites of Cyclops strenuus abyssorum Sars. Larger copepodites and adults of C. s. abyssorum and Diaptomus gracilis Sars were consumed as larvae grew and cladocerans were also eaten by larvae >11 mm. The diet of juveniles included a wider range of cladocerans and benthic invertebrates. Cannibalism was not observed among 0+ perch.
• 3 The range of food items taken by larvae increased with length, and the size of food particles ingested was governed by jaw gape which was linearly related to fish length.
• 4 Larvae fed during daylight hours with feeding maxima in midmorning and the evening.
• 5 The arithmetic (constant) rate of evacuation R of the digestive tract of larvae was 0.59—1.16% of body weight h^?1 and exponential R was 0.21–0.33% h^?1. Daily C_t calculated using an arithmetic model was 14.2–23.4% body weight at water temperatures of 12.5–18.2°C, and 23.9–40.3% utilizing an exponential method. For larvae of 9 mm. this was 46 μg dry weight day^?1 and 148 by an exponential model. Food conversion efficiency was 37–72% (arithmetic model) and 21–45% (exponential model).

     

Biogeographic conundrum: Why so few stream nerite species (Gastropoda: Neritidae) in Australia?

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Studies on the biology of Chaoborus flavicans (Meigen) (Diptera: Chaoboridae) in a fish-free eutrophic pond,Japan

Population dynamics of Chaoborus flavicans larvae of various instars was studied from November 1986 to December 1987 in a eutrophic, fish-free pond, Japan. First and 2nd instar larvae were observed from late April to late October, indicating a reproductive period of about half a year. C. flavicans overwintered in the 4th instar larvae. In water column samples, total density of all instars was 680–23680 m^-2, and pupal density 0–2600 m^-2; larvae of the 1st, 2nd, and 3rd instars showed 5–6 density peaks in 1987, suggesting that 5–6 generations occur during a year (peaks of the 4th instar larvae were not clear, probably due to their longer development than those of younger instars). In sediment samples, no 1st and 2nd instar larvae were found, 3rd instar larvae were found occasionally but density of the 4th instar larvae was 280–18600 m^-2, and pupal density varied between 0–502 m^-2. Fouth instar larvae accumulated in sediment in the cold season and in the water column in the warm season; high temperature and low oxygen concentration were the most important factors limiting the distribution of larvae in the sediment in summer in the NIES pond. The dry weight of total C. flavicans larvae was 0.08–4.2 g m^-2 in sediment samples and 24–599 μg l^-1 (0.10–2.40 g m^-2) in water column samples. Comparisons of maximum densities in the NIES pond in different years and in waters of different trophic status show that density is generally higher in eutrophic than in oligotrophic habitats. This revised version was published online in July 2006 with corrections to the Cover Date.     

Wood‐boring insect abundance in fire‐injured ponderosa pine

• 1 Wood‐boring larvae in the families Cerambycidae and Buprestidae are often found in high densities in burned trees after wildland fires. They play an important role in tree decomposition, often reducing the value of salvageable timber, and represent an important avian food source.
• 2 Three forest areas that experienced wildfires 1–3 years previously were surveyed during the summer of 2004. Ponderosa pine trees with green, scorched and consumed needles were examined for wood borer occurrence. Within each of the three needle damage categories, the mean wood borer incidence was similar between different age fires. Trees with scorched or consumed needles had significantly more wood borers than trees with green needles.
• 3 Larvae collected from under tree bark were identified to family; when possible, cerambycids were identified further to Acanthocinus spp., Monochamus sp., Rhagium inquisitor (L.) and Stictoleptura canadensis (LeConte), and buprestids were identified to Chalcophora spp. and Chrysobothris sp.
• 4 Classification tree models showed that the estimated probability of tree infestation by wood borers varied among needle damage categories. For trees with green needles, tree injury variables of high bole char height and phloem discolouration were important predictive variables. In trees where needles were consumed, tree size variables of diameter at breast height and tree height were important predictive variables.
• 5 More than half the dead trees examined were infested with wood borers, indicating that infestation of fire‐killed ponderosa pine may represent an important food source for species such as woodpeckers and a potential problem for the utilization of infested trees.

     

Life cycle and growth ofTrichostegia minor (Curtis) in temporary woodland pools (Trichoptera: Phryganeidae)

Larvae of the caddisTrichostegia minor (Curtis) were collected from four woodland pools in The Netherlands, three of which are temporary, from August 1986 till June 1987. Eggs and larvae of this species proved to be very well adapted to drought, freezing, strongly fluctuating pH and alkalinity levels and prolonged oxygen deficit. The life cycle ofT.minor in a small woodland marsh overgrown byCalla palustris took one year. Adult flight period started at the end of May. Oviposition took place independent of water. Hatching of the eggs started in September and was probably induced by immersion. During the larval stage from September until May, 5 instars could be distinguished by the size of the head capsule. Growth of instars I, II and III during autumn was moderate. Most larvae overwintered as instar III or IV. Possibly there was a larval diapause during winter. In spring rapid growth to instar V took place prior to pupation. Growth rate, expressed as the increase of mean individual dry weight was highest from March to April (2.05±0.75% DW.m^–2.d^–1). In extremely shallow water growth in spring was initially more rapid compared to growth in deeper water. During winter the growth rate decreased to 0.038±0.071% DW.m^–2.d^–1. Net annual production based on the changes of momentary biomass was 183.2±31.7 mg DW.m^–2.y^–1 or 177.2±31.3 mg AFDW.m^–2.y^–1. Production loss during the winter season was 75.1±10.8 mg DW.m^–2.y^–1 or 72.3±10.6 mg AFDW.m^–2.y^–1.     

High temperature tolerance and thermal plasticity in emerald ash borer Agrilus planipennis

• 1 The emerald ash borer Agrilus planipennis (Coleoptera: Buprestidae) (EAB), an invasive wood‐boring beetle, has recently caused significant losses of native ash (Fraxinus spp.) trees in North America. Movement of wood products has facilitated EAB spread, and heat sanitation of wooden materials according to International Standards for Phytosanitary Measures No. 15 (ISPM 15) is used to prevent this.
• 2 In the present study, we assessed the thermal conditions experienced during a typical heat‐treatment at a facility using protocols for pallet wood treatment under policy PI‐07, as implemented in Canada. The basal high temperature tolerance of EAB larvae and pupae was determined, and the observed heating rates were used to investigate whether the heat shock response and expression of heat shock proteins occurred in fourth‐instar larvae.
• 3 The temperature regime during heat treatment greatly exceeded the ISPM 15 requirements of 56 °C for 30 min. Emerald ash borer larvae were highly tolerant of elevated temperatures, with some instars surviving exposure to 53 °C without any heat pre‐treatments. High temperature survival was increased by either slow warming or pre‐exposure to elevated temperatures and a recovery regime that was accompanied by up‐regulated hsp70 expression under some of these conditions.
• 4 Because EAB is highly heat tolerant and exhibits a fully functional heat shock response, we conclude that greater survival than measured in vitro is possible under industry treatment conditions (with the larvae still embedded in the wood). We propose that the phenotypic plasticity of EAB may lead to high temperature tolerance very close to conditions experienced in an ISPM 15 standard treatment.

     

Swimming Behaviour of <Emphasis Type="Italic">Chironomus acerbiphilus</Emphasis> Larvae in Lake Katanuma

We conducted a seasonal survey of the swimming behaviour of Chironomus acerbiphilus larvae in volcanic Lake Katanuma from April 1998 to December 2001. Swimming C. acerbiphilus density was much higher than other chironomid species in lakes. All C. acerbiphilus larvae (1st through 4th instars) swam, but the earlier instars (especially the 1st) had the greatest densities and fluctuations. First instars were never found in the benthic population. This result indicates that the 1st-instar larvae are planktonic. Low water temperature (below about 10 °C) resulted in the seasonal disappearance of swimming chironomid larvae. Chemical factors – oxygen depletion or presence of hydrogen sulfide – also restricted the distribution of swimming and benthic larvae. Larvae were distributed only in the oxygen-rich part of the lake bottom and swam only in the oxygen-rich layer of the water column. The density of older swimming C. acerbiphilus (3rd and 4th instars) tended to increase with increasing benthic larval densities. The chemical stress of oxygen depletion or presence of hydrogen sulfide during holomixis within and after the stratification period leads to conspicuous swimming behaviour of benthic C. acerbiphilus larvae. Almost all C. acerbiphilus larvae died on this occasion.     

Maternal behaviours may explain riffle‐scale variations in some stream insect populations

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Wood borer performance and wood characteristics of drought‐stressed Scots pine seedlings

Four‐year‐old Scots pine [Pinus sylvestris L. (Pinaceae)] seedlings were exposed to medium and severe drought stress for two consecutive years. The anatomical properties of drought‐stressed Scots pine wood and their impact on the performance of destructive wood boring early instars of Hylotrupes bajulus L. (Coleoptera: Cerambycidae) were studied. Drought stress significantly decreased diameter of earlywood tracheids in both growing years and diameter of latewood tracheids after the second growing season only. Cell lumen area was significantly decreased by both medium and severe drought stress compared to well‐watered controls. In addition, area of cell lumen was significantly smaller in severe drought than in medium drought treatment. The drought stress marginally increased the number of resin canals in the wood, but did not affect the size of resin canals either in wood or bark. The relative growth rate of xylophagous H. bajulus neonatal larvae was not significantly affected by drought stress during the 106‐day feeding period on Scots pine wood blocks. The results show that although water availability was an important factor affecting the development and anatomy of wood cells, observed changes in wood characteristics did not affect the performance of early instars feeding on wood processed from drought‐stressed young Scots pine seedlings.     

Biofilm development and invertebrate colonization of wood in four New Zealand streams of contrasting pH

1. Biofilm development and activity on wood substrata (Nothofagus menziesii) were examined at four forested sites in a South Island, New Zealand, river catchment over a period of 6 months. Two of the sites had brown waters and mean pH of 3.7 and 4.5, whereas the other two had clear waters and mean pH of 6.3 and 6.8. 2. Fungi and other filamentous heterotrophs were the dominant colonizers of wood at all sites; few algal cells were present. Incorporation of ¹⁴C-glucose by biofilms was greatest in all four streams after 3 months, whereas endocellulase activity fluctuated over time and temporal patterns differed among streams. 3. No clear relationship was found between the incorporation of ¹⁴C-glucose or endocellulase activity of biofilms and pH, although at one near-neutral pH site ¹⁴C-glucose uptake increased in response to nutrient (N + P) additions. 4. After 6 months, incorporation of ¹⁴C-glucose and endocellulase activity of biofilms on Pinus radiata dowels buried vertically in the stream beds did not differ at depths of 3–9 cm and 19–25 cm in each stream. 5. Radiotracer experiments with a grazing amphipod (Paraleptamphopus sp.) demonstrated that biofilms on wood from all four sites could be ingested and at least partially assimilated. Chironomid larvae and harpacricoid copepods were the most abundant invertebrates colonizing wood substrata at all sites. Different chironomid species dominated at acidic and near-neutral pH sites. 6. Overall, our findings provide little support for the hypothesis that microbial activity on organic substrata is necessarily lower in streams of low pH.     

Contrasting diel activity and feeding patterns of four instars of Rhyacophila dorsalis (Trichoptera)

1. Ontogenetic shifts in prey choice and predator behaviour can affect food‐web structure. Therefore, it is important to establish if the diet and feeding activity differ between life‐stages of the same species. This hypothesis was tested for second, third, fourth and fifth larval instars of Rhyacophila dorsalis by comparing their diel activity and feeding patterns. Second to fifth instars collected from two streams were used either for gut analyses or for observations of their activity and feeding patterns in three stream tanks. Food was provided in excess; being organisms living in bryophytes on top of a large stone in each tank, augmented by different‐sized larvae of Ephemeroptera, Simuliidae and Chironomidae. As few first instars for gut analyses were found in the field, the diet of first instars reared in the laboratory was also studied. 2. Larvae for gut analyses were taken 1 h before dusk or dawn (n = 50 larvae per instar for each day or night sample). First and second instars fed on the smaller food items with no significant day‐night differences in diet. Gut contents indicated a progressive trend from feeding chiefly at night in third instars to almost exclusively at night in fifth instars. Fourth and fifth instars fed on the larger food items, whilst the diet of the third instar larvae overlapped with that of both the earlier and later instars. 3. Diel activity patterns of single larvae differed between instars but not within each instar (n = 20 larvae per instar). Second instars were active throughout the 24 h, with peaks at dusk, around midnight, dawn and around midday. A similar pattern was shown by third instars but the peak of activity at midday was less than the other three peaks. Prey were captured only during these peaks for both instars. Fourth and fifth instars were most active, and fed only, at night. They used an ambush strategy to capture more active prey at dusk and dawn (e.g. Baetis, Gammarus), and a searching strategy to capture more sedentary prey during the night (e.g. chironomids, simuliids). These experiments provided support for the hypothesis under test. If competition and/or interference occur between instars, then it could be reduced between earlier and later instars because of differences in their diet and diel pattern of feeding activity.