Seasonal nitrogen dynamics of Miscanthus×giganteus and Panicum virgatum

There is a tradeoff to consider when harvesting perennial biomass crops: harvest too late and yield declines, harvest too early and risk higher mineral contents, particularly nitrogen (N). Allowing the crop to completely senesce and recycle nutrients has several advantages, including improved feedstock quality and reduced fertilizer requirements, but it comes at a risk, particularly in temperate climates where snow and ice can reduce or destroy harvestable biomass. The effect of harvest time on the N concentration ([N]) and biomass of Panicum virgatum and Miscanthus × giganteus was evaluated at three sites in Illinois over two years. In both species [N] of standing biomass significantly declined with time ( P <0.0001). Interestingly, there was no significant interaction effect of species and sample date on [N] ( P =0.2888), but there was a highly significant interaction effect on the total N in standing biomass ( P <0.0001). The amount of standing N was directly related to biomass yield. Seasonal changes in standing N differed among locations, suggesting that harvest time recommendations for N management depend on location. P. virgatum would have potentially removed as much as 187 kg N ha⁻¹ if harvested green, and as little as 5 kg N ha⁻¹ if harvested in late winter. Because of higher biomass yields, M . × giganteus standing N ranged from 379 kg N ha⁻¹ in June to <17 kg N ha⁻¹ in February. Importantly, there was little overall change in [N] between an early winter (December) harvest and a late winter (February/March) harvest, indicating the benefits of N cycling in the system can be realized by end of the growing season and thus, at least from an N economy perspective, there is no reason to risk yield losses by delaying harvest over the winter.     

Biomass and carbon accumulation in a fire chronosequence of a seasonally dry tropical forest

Seasonally dry tropical forests (SDTF) are a widely distributed vegetation type in the tropics, characterized by seasonal rainfall with several months of drought when they are subject to fire. This study is one of the first attempts to quantify above- and belowground biomass (AGB and BGB) and above- and belowground carbon (AGC and BGC) pools to calculate their recovery after fire, using a chronosequence approach (six forests that ranged form 1 to 29 years after fire and mature forest). We quantified AGB and AGC pools of trees, lianas, palms, and seedlings, and BGB and BGC pools (Oi, Oe, Oa soil horizons, and fine roots). Total AGC ranged from 0.05 to nearly 72 Mg C ha⁻¹, BGC from 21.6 to nearly 85 Mg C ha⁻¹, and total ecosystem carbon from 21.7 to 153.5 Mg C ha⁻¹; all these pools increased with forest age. Nearly 50% of the total ecosystem carbon was stored in the Oa horizon of mature forests, and up to 90% was stored in the Oa-horizon of early successional SDTF stands. The soils were shallow with a depth of <20 cm at the study site. To recover values similar to mature forests, BGC and BGB required <19 years with accumulation rates greater than 20 Mg C ha⁻¹ yr⁻¹, while AGB required 80 years with accumulation rates nearly 2.5 Mg C ha⁻¹ yr⁻¹. Total ecosystem biomass and carbon required 70 and 50 years, respectively, to recover values similar to mature forests. When belowground pools are not included in the calculation of total ecosystem biomass or carbon recovery, we estimated an overestimation of 10 and 30 years, respectively.     

Carbon mitigation by the energy crop, Miscanthus

Biomass crops mitigate carbon emissions by both fossil fuel substitution and sequestration of carbon in the soil. We grew Miscanthus x giganteus for 16 years at a site in southern Ireland to (i) compare methods of propagation, (ii) compare response to fertilizer application and quantify nutrient offtakes, (iii) measure long-term annual biomass yields, (iv) estimate carbon sequestration to the soil and (v) quantify the carbon mitigation by the crop. There was no significant difference in the yield between plants established from rhizome cuttings or by micro-propagation. Annual off-takes of N and P were easily met by soil reserves, but soil K reserves were low in unfertilized plots. Potassium deficiency was associated with lower harvestable yield. Yields increased for 5 years following establishment but after 10 years showed some decline which could not be accounted for by the climate driven growth model MISCANMOD. Measured yields were normalized to estimate both autumn (at first frost) and spring harvests (15 March of the subsequent year). Average autumn and spring yields over the 15 harvest years were 13.4±1.1 and 9.0±0.7 t DW ha⁻¹ yr⁻¹ respectively. Below ground biomass in February 2002 was 20.6±4.6 t DW ha⁻¹. Miscanthus derived soil organic carbon sequestration detected by a change in ¹³C signal was 8.9±2.4 t C ha⁻¹ over 15 years. We estimate total carbon mitigation by this crop over 15 years ranged from 5.2 to 7.2 t C ha⁻¹ yr⁻¹ depending on the harvest time.     

Trade-off in oviposition strategy: choosing poor quality host plants reduces mortality from natural enemies for a salt marsh planthopper

Abstract.  1. Both host plant nutrition and mortality from natural enemies have been predicted to significantly impact host plant selection and oviposition behaviour of phytophagous insects. It is unclear, however, if oviposition decisions maximise fitness.
2. This study examined whether the salt marsh planthopper Pissonotus quadripustulatus prefers higher quality host plants for oviposition, and if oviposition decisions are made so as to minimise mortality at the egg stage.
3. A controlled laboratory experiment and 4 years of field data were used to assess the rates of planthopper oviposition on higher quality 'green' and lower quality 'woody' stems of the host plant Borrichia frutescens . The numbers and percentages of healthy eggs and eggs that were killed by parasitoids or the host plant were recorded.
4. In all years, including the laboratory experiment, Pissonotus planthoppers laid more eggs on lower quality woody stems than on higher quality green stems. While host plant related egg mortality was higher in woody stems, the percentage of eggs parasitised was much greater in green stems. This resulted in a lower total mortality of eggs on woody stems.
5. The results of this study demonstrate that, although Pissonotus prefers lower quality host plants for oviposition, this actually increases fitness. These data seem to support the enemy free space hypothesis, and suggest that for phytophagous insects that experience the majority of mortality in the egg stage, oviposition choices may be made such that mortality is minimised.     

Influence of host plant vs. natural enemies on the spatial distribution of a pine sawfly, Neodiprion autumnalis

Abstract .1. The pine sawfly, Neodiprion autumnalis , infests ponderosa pine, Pinus ponderosa , growing at low densities near the bottom of an altitudinal gradient in Arizona, U.S.A. The relative importance of host-plant quality vs. natural-enemy effects in determining the spatial distribution of this sawfly was examined over a 3-year period.
2. Field and laboratory bioassays were conducted on all life stages of N. autumnalis at two forest stand densities (high ≥ 23 m² ha^–1, low ≤ 7 m² ha^–1) and at two elevations (bottom slope = 2390 m, top slope = 2540 m). These experiments were used for constructing life tables of N. autumnalis that compared the effects of host-plant quality on oviposition preference and progeny performance with the effects of natural enemies at different tree densities and elevations.
3. Life-table analyses determined that mortality attributed to host-plant effects during the egg and larval stages had the largest impact on fitness between tree densities and elevations.
4. Natural enemies caused a significant reduction in progeny survival, but their effects were similar across all tree densities and elevations during egg and larval life stages. However, cocoon-stage survival did vary between tree densities and elevations due to natural-enemy effects.
5. It was concluded that the observed oviposition preference for, and higher progeny performance on, trees at low densities and bottom slope elevations were caused primarily by host-plant effects.
6. These results further the argument that heterogeneity at the resource level (i.e. bottom-up forces) determines potential outcomes of multitrophic level interactions.     

Fish production in the Jarama River, Central Spain

Fish production was estimated at three sites on the Jarama River, a small, typical upland river in Central Spain. Population estimates were made by the successive removal method of electrofishing. The same six species were recorded at each of the sites: Chondrostoma polylepis, Barbus barbus bocagei, Leuciscus cephalus pyraenaicus, Salmo trutta m. fario, Gobio gobio and Cobitis paludicola , with the first three species always dominant. Density, biomass, production (assuming that N_o is the total number of eggs spawned), and available production were, respectively: 13502-85776 ind. ha⁻¹, 178.6–221.3 kg ha⁻¹, 221.7–583.6 kg ha⁻¹ yr⁻¹, 118.1–271.9 kg ha⁻¹ yr⁻¹. Production estimates based on mortality curves were 7.9–19.5% (mean: 13.7) lower than those based on estimated from the number of eggs laid. Production per unit of area was highest at the widest and deepest site. Brown trout production contributed only 2–4% of the total production for all sites.     

Development, growth, and egg production of Ageneotettix deorum (Orthoptera: Acrididae) in response to spider predation risk and elevated resource quality

Abstract.  1. Predation risk to insects is often size- or stage-selective and usually decreases as prey grow. Any factor, such as food quality, that accelerates developmental and growth rates is likely to reduce the period over which prey are susceptible to size-dependent predation.
2. Using field experiments, several hypotheses that assess growth, development, and egg production rates of the rangeland grasshopper Ageneotettix deorum (Scudder) were tested in response to combinations of food quality and predation risk from wolf spiders to investigate performance variation manifested through a behaviourally mediated path affecting food ingestion rates.
3. Grasshoppers with nutritionally superior food completed development ≈ 8–18% faster and grew 15–45% larger in the absence of spiders, in comparison with those subjected to low quality food exposed to spider predators. Growth and development did not differ for grasshoppers feeding on high quality food when predators were present in comparison with lower quality food unimpeded by predators. Responses indicated a compensatory relationship between resource quality and predation risk.
4. Surviving grasshoppers produced fewer eggs compared with individuals not exposed to spiders. Because no differences were found in daily egg production rate regardless of predation treatment, lower egg production was attributed to delayed age of first reproduction. Results compare favourably with responses observed in natural populations.
5. Risk of predation from spiders greatly reduced growth, development, and ultimately egg production. Increased food quality counteracts the impact of predation risk on grasshoppers through compensatory responses, suggesting that bottom-up factors mediate effects of spiders.     

A study of the dynamics of a natural population of the freshwater crayfish, Austropotamobius pallipes

SUMMARY. 1. Population size, survival and recruitment were estimated from mark—recapture data at frequent intervals.
2. Two types of mortality were identified: (1) moult-associated—an intense but short duration mortality, and (2) overwinter mortality, which operated at a lower estimated weekly rate but over some 30 weeks.
3. Population size reached a mid-summer peak as recruitment into the catchable size classes occurred. Thereafter numbers declined slowly to the end of the growth season, followed by an overwinter decline of some 50% of total.
4. Annual production was estimated at 196.5 kg wet wt ha⁻¹ in 1977, 116.8 kg wet wt ha⁻¹ in 1978 and 87.79 kg wet wt ha⁻¹ in 1979.     

Fish production in Oued Sebaou, a seasonal river in North Algeria

Fish were sampled by electrofishing at three contiguous sites 26 km from the source of Oued Sebaou, a seasonal river in North Algeria. River discharge ranges from 300 m³s⁻¹ in the peak of the rainy season to 0.1 m³s⁻¹ at the end of the dry season. Extensive gravel extraction from the river bed takes place during the dry season.
Two species were recorded at each site: Barbus callensis , which is heavily fished locally, and Anguilla anguilla . The three sample maximum likelihood Zippin method was used to estimate mean density and standing crops, which were: barbel, 452 fish ha⁻¹, 5.33 kg ha⁻¹; eels, 149 fish ha⁻¹, 3.04 kg ha⁻¹. Estimated average biomass and production were: barbel, 2.31 kg ha⁻¹, 4.27 kg ha⁻¹ year⁻¹; eels, 1.51 kg ha⁻¹, 2.60 kg ha⁻¹ year⁻¹.     

Soil seed bank dynamics of fire-prone wooded grassland, woodland and dry forest ecosystems in Ethiopia

In order to reveal the role of soil seed banks in vegetation recovery after fire in savanna, the spatial distribution and temporal changes in the soil seed banks of regularly burning savanna in Gambella, western Ethiopia, was studied. The seedling emergence technique was employed to determine the species composition and density of the soil seed bank of six sites ranging in fire severity from wooded grassland with frequent fires over woodland with intermediate fire frequency to forest with absence of fires. Species composition and density of seeds in the soil were compared between seasons, depths and sites with different types of standing vegetation. Fourteen plant species were recorded in the soil seed bank from the grassland and woodland sites and 6 from the dry forests; 60 % of the taxa in the soil seed bank were annuals and 40 % were perennials. The soil seed banks were largely dominated by graminoids and 48–97 % of the soil seed bank in the grasslands and woodlands was of a single grass species, Hyparrhenia confinis , which was absent from the dry forests. The soil seed pools ranged from less than 100 to 4700 seeds per m² depending upon the season. The soil seed bank of graminoids was nearly empty after the onset of the rainy season whereas seeds of broadleaved herbs and woody species able to germinate were still found after this time. Floristic composition, representation of life forms and density of seeds in soil did not correspond closely with that of the standing vegetation, but within graminoids there was a strong similarity between the soil seed bank and the standing vegetation. The current fire regime of Ethiopian savanna woodlands appears to maintain the dominance of graminoids over broadleaved herbs and woody plants both as seeds in the soil and in the standing vegetation.     

Terrestrial export of highly bioavailable carbon from small boreal catchments in spring floods

1. We assessed the terrestrial export of organic carbon, which effectively supported aquatic bacterial production (BP), from small boreal catchments during spring flood. We analysed stream runoff from nine small catchments with different proportions of peat mires and coniferous forests by monitoring the dissolved organic carbon (DOC) flux in combination with conducting bacterial bioassays.
2. Multiple linear regression analysis showed that BP during 7-day-dark bioassays (BP₇; μg C L⁻¹day⁻¹) was explained by both the quantity and quality (low-molecular weight fractions) of the DOC. BP₇ can be used as a measure of export of terrestrial organic carbon that is highly bioavailable.
3. Total export of DOC during spring flood from the different catchments ranged from 20 to 27 kg ha⁻¹ and was negatively correlated to forest cover (%). However, the export of BP₇ carbon was positively correlated to forest cover and varied from about 0.1 kg ha⁻¹ in mire-dominated streams to about 0.2 kg ha⁻¹ in forest-dominated streams.
4. The high bioavailability of forest carbon suggests that forests are the main contributors of BP-supporting carbon in boreal streams although mires have higher area-specific export of DOC.     

Synchrony in small mammal populations of montane forest patches in southern India

1. Small mammal populations were studied in montane evergreen forests in the Nilgiris, southern India, using live-trapping from January 1994 to September 1996. Two sites were selected, one with a single large forest patch and adjoining plantations, and the other with several small forest patches separated by grassland.
2. Nine species were recorded, of which eight were trapped in the forest patches, two in the grasslands and five in plantations. Rattus rattus was the most abundant species (2–36 individuals ha⁻¹) followed by Suncus montanus (0–11 individuals ha⁻¹). Densities of seven other species recorded were very low.
3. Synchrony in various population variables – density, biomass, mean weight, density of adults and adult females, and proportion of adults, adult females and sub-adults – was examined for Rattus rattus populations in the forest patches and plantations. Density and biomass were studied in seven other species recorded at these sites. Spearman's rank order correlation was used as a measure of synchrony between the population variables.
4. Within-site synchrony was higher than between-site synchrony in population characters. Synchrony was also higher between plots within the unfragmented site than they were between plots in the fragmented site. Relatively high synchrony in proportions of adults, adult females and juveniles in the forest patches implied that breeding is probably influenced by climate and food availability, which are seasonal in this habitat.
5. Given the small patch sizes (≈1–10 ha) and low population sizes, asynchrony is likely to be an outcome of demographic and environmental stochasticity, and low dispersal rates may impede establishment of synchrony.     

The effect of egg load on readiness to accept a low-quality host plant is weak and age dependent in a geometrid moth

Abstract.  1. Most empirical and theoretical studies treat egg load as a major determinant of insect oviposition behaviour, and predict a positive link between egg load and readiness to oviposit.
2. In the present study, the correlation between egg load and readiness to lay eggs on a low-ranked host was examined in wild-caught moths, Scotopteryx chenopodiata L. (Lepidoptera: Geometridae).
3. No overall effect of egg load on oviposition could be detected. However, there was a significant interaction between egg load and wing wear: in only one wing-wear class, out of four, high egg load tended to promote oviposition, while in the three other classes the effect was insignificant.
4. The results suggest that the effect of egg load on oviposition may depend on the age of the ovipositing insect.     

Meeting US biofuel goals with less land: the potential of Miscanthus

Biofuels from crops are emerging as a Jekyll & Hyde – promoted by some as a means to offset fossil fuel emissions, denigrated by others as lacking sustainability and taking land from food crops. It is frequently asserted that plants convert only 0.1% of solar energy into biomass, therefore requiring unacceptable amounts of land for production of fuel feedstocks. The C₄ perennial grass Miscanthus × giganteus has proved a promising biomass crop in Europe, while switchgrass ( Panicum virgatum ) has been tested at several locations in N. America. Here, replicated side-by-side trials of these two crops were established for the first time along a latitudinal gradient in Illinois. Over 3 years of trials, Miscanthus × giganteus achieved average annual conversion efficiencies into harvestable biomass of 1.0% (30 t ha⁻¹) and a maximum of 2.0% (61 t ha⁻¹), with minimal agricultural inputs. The regionally adapted switchgrass variety Cave-in-Rock achieved somewhat lower yields, averaging 10 t ha⁻¹. Given that there has been little attempt to improve the agronomy and genetics of these grasses compared with the major grain crops, these efficiencies are the minimum of what may be achieved. At this 1.0% efficiency, 12 million hectares, or 9.3% of current US cropland, would be sufficient to provide 133 × 10⁹ L of ethanol, enough to offset one-fifth of the current US gasoline use. In contrast, maize grain from the same area of land would only provide 49 × 10⁹ L, while requiring much higher nitrogen and fossil energy inputs in its cultivation.     

Influence of egg size differences within egg clutches on larval parameters in nine libellulid species (Odonata)

Abstract.  1. In libellulids, egg size differs between species and populations. There are also size differences within egg clutches of individual females.
2. Past experiments suggest that there are two different types of egg clutches in libellulids. Egg size decreases significantly during oviposition in species that perform non-contact guarding during oviposition. In contrast, in species ovipositing in tandem, egg size is randomly distributed.
3. This study deals with the possible consequences of egg size variation within the different egg clutch types. The study examined whether there is a correlation between egg development time, offspring sex or larval size and egg size.
4. The current experiments were conducted in Namibia and Germany. Five non-contact guarding and four tandem guarding libellulid species were used.
5. In some species larger eggs needed more time to develop, in some species no correlation between egg size and egg development time could be found, whereas in other species larger eggs developed faster.
6. The sex ratio was biased towards females in Leucorrhinia dubia and in Sympetrum striolatum and egg size was not associated with gender.
7. In both egg clutch types larger eggs resulted in larger larvae. In this study, evidence was found that the effects of egg size diminished with progressing larval development under good conditions. However, it is possible that the effects may have a greater influence under harsh circumstances.     

Inactivation of Salmonella enterica serovar enteritidis in shell eggs by sequential application of heat and ozone

Aims:  To assess the contribution of ozone to lethality of Salmonella enterica serovar Enteritidis in experimentally inoculated whole shell eggs that are sequentially treated with heat and gaseous ozone in pilot-scale equipment.
Methods and Results:  Whole shell eggs were inoculated with small populations of Salmonella Enteritidis (8·5 × 10⁴–2·4 × 10⁵ CFU per egg) near the egg vitelline membrane. Eggs were subjected to immersion heating (57°C for 21 min), ozone treatment (vacuum at 67·5 kPa, followed by ozonation at a maximum concentration of approx. 140 g ozone m⁻³ and 184–198 kPa for 40 min) or a combination of both treatments. Survivors were detected after an enrichment process or enumerated using modified most probable number technique. Ozone, heat and combination treatments inactivated 0·11, 3·1 and 4·2 log Salmonella Enteritidis per egg, respectively.
Conclusions:  Sequential application of heat and gaseous ozone was significantly more effective than either heat or ozone alone. The demonstrated synergy between these treatment steps should produce safer shell eggs than the heat treatment alone.
Significance and Impact of the Study:  Shell eggs are the most common vehicle for human infection by Salmonella Enteritidis. Many cases of egg-related salmonellosis are reported annually despite efforts to reduce contamination, including thermal pasteurization of shell eggs and egg products. Treatment with ozone-based combination should produce shell eggs safer than those treated with heat alone.     

Gallery construction and oviposition of the bark beetle Ips typographus (Coleoptera: Scolytidae) at different breeding densities

Abstract. 1. The bole of a Norway spruce, Picea abies (L.) Karst., was cut in 25 cm sections and infested in the laboratory with 2 or 8 females/dm² of Ips typographus (L.) and an equal number of males.
2. One log of each beetle density was dissected every second day, beginning 2 days after female introduction.
3. The oviposition rate was highest at the beginning of the breeding period for both densities and then declined, especially at the high density. Residence time is a useful approximation for the number of laid eggs in the beginning of the oviposition period or when the density is low.
4. Final gallery length and egg number per gallery were on average significantly smaller at density 8 than at density 2.
5. Linear relationships between total gallery length and number of egg niches were found to be similar for the two densities.
6. An estimate of the egg to adult mortality at different densities was found by using data from a field study that related gallery length to breeding density in attacked living trees, together with published relationships between density and egg niches or emergence of new adults.
7. It was concluded that both decreased oviposition rate and earlier re-emergence at higher densities contribute to a declining oviposition with increasing density. The final offspring production is futher reduced by lower survival due to larval competition as density increases.     

The global relationship between forest productivity and biomass

Aim  We aim to determine the empirical relationship between above-ground forest productivity and biomass. There are theoretical reasons to assume a relationship between forest structure and function, as both may be influenced by similar ecological factors such as moisture supply. Also, dynamic global vegetation model simulations imply that any increase in forest productivity driven by climate change will result in increases in biomass and therefore carbon storage. However, few studies have explored the strength and form of the relationship between forest productivity and biomass, whether in space or time.
Location Global scale.
Methods  We collated a large data set of above-ground biomass (AGB) and above-ground net primary productivity (ANPP) and tested the extent to which spatial variation in forest biomass across the Earth can be predicted from forest productivity.
Results  The global ANPP–AGB relationship differs fundamentally from the strongly positive, linear relationship reported in earlier analyses, which mostly lacked tropical sites. AGB begins to peak at c . 15–20 Mg ha⁻¹ year⁻¹ ANPP, plateaus at ANPP > 20–25 Mg ha⁻¹ year⁻¹, and may actually decline at higher levels of production.
Main conclusions  High turnover rates in high-productivity forests may limit AGB by promoting the dominance of species with a low wood density. Predicted increases in ANPP will not necessarily favour increases in forest carbon storage, especially if changes in productivity are accompanied by compositional shifts.     

The fate of a roach Rutilus rutilus stock under an extremely strong fishing pressure and its predicted development after the cessation of mass removal

The effects of very intensive exploitation on a roach stock over 5 years are estimated. The total roach catch of the mass removal carried out with trawls was 507 metric tons (195 kg ha⁻¹, 54% of the total catch). According to the virtual population analysis the biomass of the stock has decreased from 180 kg ha⁻¹ in 1989 to c . 50 kg ha⁻¹ in 1993. The production of the stock has fallen from 64 to 33 kg ha⁻¹. The stock is dominated by younger age groups than before the mass removal. The future development of the stock is predicted by considering different levels of natural and fishing mortality. If the exploitation is stopped totally after 1993, roach biomass is predicted to double in 3 years. A constant fishing mortality rate of 0.3 (yearly catches 45–50 metric tons) would prevent the recovery of the stock, providing that 1–year–old and older roaches are exploited.     

The influence of darkness and visual contrast on oviposition by Anopheles gambiae in moist and dry substrates

Abstract.  The influence of substrate brightness (value), contrast and colour (hue) on oviposition is quantified under low night-light levels of 1.2 × 10⁻⁴ w m⁻² in choice-tests using caged Anopheles gambiae sensu stricto laboratory strains and/or gravid An. gambiae sensu lato from a natural population in Western Kenya. Egg output increases as darkness of the substrate increases against a white background. Black substrates elicit the most oviposition by both laboratory and house-collected mosquitoes, whereas white elicits the least. Contrast between patches of substrate and background also influences egg placement. In four-choice tests, a black ovipositional dish against a white floor receives many more eggs than does black–black, white–white, and white–black dish vs. floor combinations. In a similar test presenting four grey vs. black combinations of dish vs. floor, females respond similarly to grey vs. black dishes and deposit nearly as many eggs on the dry floors as on wet dishes. Illumination at 2.1 × 10⁻³ w m⁻², equivalent to late dusk and early dawn, improves discrimination between black dishes and grey floors. Under the experimental conditions, An. gambiae females respond almost equally to dishes of water placed over red, yellow, green, blue and purple paper disks matching the brightness and saturation values of their neutral (grey) counterparts. Thus, no evidence is found for wavelength discrimination on oviposition. It is concluded that visual contrast strongly influences the search patterns for prospective ovipositional sites whereas darkness of substrate and moisture are strong releasers of oviposition.