The influence of fungal food quality on the growth and fecundity of Folsomia candida (Collembola: Isotomidae)

Summary The Basidiomycete fungi Coriolus versicolor and Hypholoma fasciculare were grown in liquid media containing 2, 20, 200 and 2,000 ppm nitrogen (as asparagine) and fed to cultures of Folsomia candida. Collembola feeding on both species of fungi exhibited trends of increased moulting and egg laying rates up to 200 ppm N and an inhibition of growth and fecundity at 2,000 ppm N. The differences in moulting rates between individual treatments were small for both species of fungi and not all the pair wise comparisons of treatments were significantly different. Egg laying rates of collembola fed C. versicolor showed a highly significant response to all levels of N in the growth medium and egg production at 200 ppm N was over three times higher than at 2 ppm N. Collembola fed H. fasciculare showed a less marked fecundity response to the different nitrogen levels and egg production at 200 ppm was approximately 1.5 times higher than at 2 ppm N. Both moulting and egg laying rates were significantly affected by the species of fungus presented as food to the collembola. The patterns of growth and reproduction of starved control groups of F. candida as well as those fed the test fungi demonstrate the adaptability of this species to changes in the quality and quantity of available food.     

Collembola foraging responses to interacting fungi

Abstract 1. Temperate woodland Collembola feed predominantly on plant detritus and fungi. They exhibit preferences for different fungi reflecting resource value in terms of Collembola growth and reproduction. 2. Studies of fungal grazing by Collembola have focused on non‐interacting fungi and have used one genetic isolate as representative of an entire fungal species. This study examines the effects of fungal interactions on Collembola behaviour, and elucidates differences arising from the use of genetically distinct fungal isolates. 3. Folsomia candida were added to microcosms in which paired combinations of the fungi, Hypholoma fasciculare (four isolates), Phallus impudicus, Phanerochetae velutina and Resinicium bicolor (two isolates), were interacting. Collembola movement and mortality was recorded for 26 days. A constant preference for one fungal mycelium over the other was shown by Collembola in 12 of the 36 interactions studied. Lowest Collembola mortality was normally recorded on the preferred mycelium. 4. In 11 of the remaining 24 interactions, Collembola preference switched from one mycelium to the other; the greatest number of dead Collembola was recorded on the final preference. 5. Collembola behaviour on different fungal isolates was broadly similar, although in one species’ combination a change in fungal isolate resulted in a total reversal of Collembola preference. The implications of these results for decomposer food webs are considered.     

Diet and dominance among female vervet monkeys (Cercopithecus aethiops)

Behavioral observations of free-ranging female vervet monkeys (Cercopithecus aethiops) suggest that priority of access to food enhances individual fitness. Seventeen adult females living in two social groups were observed for 26 months in the Samburu/Isiolo Game Reserves, northern Kenya. Priority of access to feeding and sitting sites was associated with significant differences in diet and reproduction. Food species distribution altered rank-related differences in diet. Rank-related differences in diet occurred when food items were clumped in distribution, but not when they were randomly distributed. The relation of individual food items to reproduction influenced rank-related differences in reproduction. Rank was significantly correlated with birth rate when a preferred food species was clumped in distribution.     

Zooplankton interactions with toxic phytoplankton: Some implications for food web studies and algal defence strategies of feeding selectivity behaviour, toxin dilution and phytoplankton population diversity

This study focuses on the interactions between toxic phytoplankton and zooplankton grazers. The experimental conditions used are an attempt to simulate situations that have, so far, received little attention. We presume the phytoplankton community to be a set of species where a population of a toxic species is intrinsically diverse by the presence of coexisting strains with different toxic properties. The other species in the community may not always be high-quality food for herbivorous zooplankton. Zooplankton populations may have developed adaptive responses to sympatric toxic phytoplankton species. Zooplankton grazers may perform a specific feeding behaviour and its consequences on fitness will depend on the species ingested, the effect of toxins, and the presence of mechanisms of toxin dilution and compensatory feeding. Our target species are a strain of the dinoflagellate Alexandrium minutum and a sympatric population of the copepod Acartia clausi. Mixed diets were used with two kinds of A. minutum cells: non-toxic and toxic. The flagellate Rhodomonas baltica and the non-toxic dinoflagellate Alexandrium tamarense were added as accompanying species. The effect of each alga was studied in separate diets. The toxic A. minutum cells were shown to have negative effects on egg production, hatching success and total reproductive output, while, in terms of its effect on fitness, the non-toxic A. minutum was the best quality food offered. R. baltica and A. tamarense were in intermediate positions. In the mixed diets, copepods showed a strong preference for toxic A. minutum cells and a weaker one for A. tamarense cells, while non-toxic A. minutum was slightly negatively selected and R. baltica strongly negatively selected. Although the level of toxins accumulated by copepods was very similar, in both the diet with only toxic A. minutum cells and in the mixed diet, the negative effects on fitness in the mixed diet could be offset by toxin dilution mechanisms. The implications of these findings are the fact that mesozooplankton may not play an important role in phytoplankton blooms development. Phytoplankton endotoxin production does not seem to be an evolutionary stable strategy as a defence against some herbivores.     

The value of three cereal aphid species as food for a generalist predator

The value of the cereal aphid species Metopolophium dirhodum (Wlk.), Sitobion avenae (F.) and Rhopalosiphum padi (L.) as prey for the linyphiid spider Erigone atra (Bl.) was assessed. Fecundity of females was determined for spiders fed on eight experimental diets: three single‐species aphid diets, a mixed diet of all three aphid species, three mixed diets with each aphid species in combination with fruit flies Drosophila melanogaster (Meig.), and pure D. melanogaster as a high quality comparison diet. The development and survival of first‐instar juveniles fed on three diets of single aphid species, and on a diet of Collembola were compared with those subjected to starvation. Prey value for adult females was assessed by egg production, hatching success and offspring size. In pure diets all three aphid species were of low value to the spiders, causing a rapid decline in egg production and supporting no growth of significance of first‐instar juveniles. No difference in value of aphid species of single‐species aphid diets was found in the fecundity experiment, while a ranking of aphid species of M. dirhodum > R. padi > S. avenae was revealed in the survivorship experiment. A mixed‐aphid diet was not found to be advantageous compared with single‐species aphid diets, and no advantage of including aphids in mixed diets with fruit flies was found. Metopolophium dirhodum and R. padi were neutral in mixed diets, while a diet of S. avenae and fruit flies caused reduced egg production compared with the pure diet of fruit flies, revealing a toxic effect of S. avenae on the spider. The value‐ranking of aphid species in mixed diets was similar to that of single‐species diets. A similar ranking of aphid species was found for different fitness parameters (fecundity of adult females and development of juveniles). A ranking of aphids by offspring size of mothers on aphid‐only diets was S. avenae > M. dirhodum > R. padi. All aphid‐fruit fly diets resulted in larger offspring than a diet of only D. melanogaster, with the overall largest offspring being produced on the diet of M. dirhodum and fruit flies.     

Being a generalist herbivore in a diverse world: how do diets from different grasslands influence food plant selection and fitness of the grasshopper Chorthippus parallelus?

1. Generalist insect herbivores occupy a variety of habitats that differ in food plant composition. Dietary mixing has been proposed as a possibility for generalists to overcome nutritional deficiencies of single plant species, but only a few studies have investigated herbivore feeding and fitness for diets that resemble natural scenarios. We studied feeding behaviour, survival, and reproduction of the generalist grasshopper Chorthippus parallelus raised on food plants of four typical habitats. 2. Grasshopper diet consisted of grasses (92.5%), legumes (6.7%) and, in small quantities, other forbs (0.8%). Diet selection differed between the four food plant mixtures, and depended on grasshopper sex and developmental stage. There was no correlation between the relative abundance of plant species in the field and the fraction of these species in the grasshopper diet. 3. Grasshoppers survived on average for 40.4 ± 1.0 days before maturity, grew 106.8 mg until maturity moult, and females laid 4.1 ± 0.4 egg pods, each of which contained 8.5 ± 0.4 eggs. However, despite the differences in feeding behaviour, grasshopper fitness was the same in all of the four food plant mixtures. While the digestibility of ingested food was similar in the four different treatments, indices indicated differences in the conversion efficiency to body mass. 4. Our results show that C. parallelus is a plastic feeder with no fixed preferences in diet composition. The results emphasise that generalist herbivores can counteract putative quality deficiencies of single food plants by selective dietary mixing.     

Invertebrate biodiversity affects predator fitness and hence potential to control pests in crops

Natural enemies that control pests usually allow farmers to avoid, or reduce, the use of pesticides. However, modern farming practices, that maximize yields, are resulting in loss of biodiversity, particularly prey diversity. Does this matter? Pests continue to thrive, and without alternative prey the predators should, perforce, concentrate their attentions upon the pests.We showed that a diverse diet significantly enhances predator fecundity and survival. Experiments were conducted using common generalist predators found in arable fields in Europe, the carabid beetle Pterostichus melanarius (Coleoptera: Carabidae) and the linyphiid spider Erigone atra (Araneae: Linyphiidae). We tested the hypothesis that mixed species diets were optimal, compared with restricted diets, with respect to parameters such as predator weights, egg weights, numbers of eggs laid, egg development times, egg hatching rates and predator survival. In carabids, an exclusive earthworm diet was as good as mixed diets containing earthworms for egg production and hatching, but less good than such mixed diets for increase in beetle mass and sustained egg laying. For spiders, aphids alone (Sitobion avenae) or with the Collembola Folsomia candida, drastically reduced survival. Aphids plus the Collembola Isotoma anglicana improved survival but only aphids with a mixed Collembola diet maximized numbers of hatching eggs.Predators offered only pests (slugs or aphids) had lowest growth rates and fecundity. We therefore demonstrated that conservation of a diversity of prey species within farmland, allowing predators to exploit a diverse diet, is essential if predators are to continue to thrive in crops and regulate agricultural pests.     

Melanin targets LC3-associated phagocytosis (LAP): A novel pathogenetic mechanism in fungal disease

Intracellular swelling of conidia of the major human airborne fungal pathogen Aspergillus fumigatus results in surface exposure of immunostimulatory pathogen-associated molecular patterns (PAMPs) and triggers activation of a specialized autophagy pathway called LC3-associated phagocytosis (LAP) to promote fungal killing. We have recently discovered that, apart from PAMPs exposure, cell wall melanin removal during germination of A. fumigatus is a prerequisite for activation of LAP. Importantly, melanin promotes fungal pathogenicity via targeting LAP, as a melanin-deficient A. fumigatus mutant restores its virulence upon conditional inactivation of Atg5 in hematopoietic cells of mice. Mechanistically, fungal cell wall melanin selectively excludes the CYBA/p22phox subunit of NADPH oxidase from the phagosome to inhibit LAP, without interfering with signaling regulating cytokine responses. Notably, inhibition of LAP is a general property of melanin pigments, a finding with broad physiological implications.     

Effect of diet on the fecundity of three carabid beetles

Abstract In the present study, the importance of diet in terms of fecundity is compared for three species of the carabid genus Amara (Coleoptera: Carabidae), using an insect diet, two types of seed diet (Capsella bursa‐pastoris, Stellaria media) and a mixed diet. It is expected that the species of carabid studied have different food requirements for reproduction. Diet affects reproduction performance and egg production significantly. A mixed diet and both single‐seed diets are suitable for reproduction in Amara aenea (DeGeer) because a higher proportion of the females reproduce and lay significantly more eggs than on a purely insect diet. Females of Amara familiaris (Duftschmid) do not reproduce unless provided with seeds of S. media. Seeds of C. bursa‐pastoris or a mixed diet are equally suitable diets for reproduction of Amara similata (Gyllenhal); a diet of insects or seeds of S. media is unsuitable. The results support the hypothesis that the species under investigation have specific food requirements, suggesting that seed feeding has evolved to different degrees in particular species: A. aenea is omnivorous, whereas A. familiaris and A. similata specialize on the seed of a particular plant species or family. This resource partitioning facilitates co‐occurrence of carabid species.     

Fruit,flies and filamentous fungi – experimental analysis of animal–microbe competition using Drosophila melanogaster and Aspergillus mould as a model system

In addition to their fundamental role in nutrient recycling, saprobiotic microorganisms may be considered as typical consumers of food‐limited ephemeral resource patches. As such, they may be engaged in inter‐specific competition with saprophagous animals feeding from the same resource. Bacteria and filamentous fungi are known to synthesise secondary metabolites, some of which are toxic and have been proposed to deter or harm animals. The microorganisms may, however, also be negatively affected if saprophagous animals do not avoid microbe‐laden resources but feed in the presence of microbial competitors. We hypothesised that filamentous fungi compete with saprophagous insects, whereby secondary metabolites provide a chemical shield against the insect competitors. For testing this, we developed a new ecological model system representing a case of animal–microbe competition between saprobiotic organisms, comprising Drosophila melanogaster and species of the fungus Aspergillus (A. nidulans, A. fumigatus, A. flavus). Infestation of Drosophila breeding substrate with proliferating fungal colonies caused graduated larval mortality that strongly depended on mould species and colony age. Confrontation with conidiospores only, did not result in significant changes in larval survival, suggesting that insect death may not be ascribed to pathogenic effects. When confronted with colonies of transgenic fungi that lack the ability to express the global secondary metabolite regulator LaeA (ΔlaeA), larval mortality was significantly reduced compared to the impact of the wild type strains. Yet, also in the ΔlaeA strains, inter‐specific variation in the influence on insect growth occurred. Competition with Drosophila larvae impaired fungal growth, however, wild type colonies of A. nidulans and A. flavus recovered more rapidly from insect competition than the corresponding ΔlaeA mutants (not in A. fumigatus). Our findings provide genetic evidence that toxic secondary metabolites synthesised by saprotrophic fungi may serve as a means to combat insect competitors. Variation in the ability of LaeA to control expression of various secondary metabolite gene clusters might explain the observed species‐specific variation in Drosophila–Aspergillus competition.     

The value of two Collembola species as food for a linyphiid spider

Egg production by the cereal spider Erigone atra was used as a fitness parameter for evaluating the food quality of two species of Collembola: Folsomia fimetaria and Isotoma anglicana. Drosophila melanogaster was used as reference prey. We tested the hypothesis that due to differences in food quality, the two Collembola species would affect the reproduction of the spider differently. The quality ranking of the prey types turned out as: I. anglicana > D. melanogaster > F. fimetaria. With F. fimetaria alone, spiders were unable to maintain reproduction. E. atra was more efficient in utilising I. anglicana and D. melanogaster. Thus, daily consumption rates of I. anglicana were lower in spite of higher egg laying rates by E. atra. A mixed diet of F. fimetaria and D. melanogaster resulted in a lower reproductive output than a pure diet of D. melanogaster, indicating a toxic element in F. fimetaria. In the mixed diet F. fimetaria had a negative influence on the consumption capacity of the spider towards D. melanogaster, while D. melanogaster had a positive influence on the consumption capacity towards F. fimetaria. It is concluded that a high abundance of I. anglicana may support a high reproductive output of E. atra, while the presence, of F. fimetaria in fields may reduce the spider's reproductive output.     

Comparison of the fecundity of three species of grain store mites on fungal diets

The egg production of isolated pairs ofAcarus siro, Glycyphagus destructor andTyrophagus longior fed on a control diet of wheatgerm and yeast was compared with that on mycelial pellets from shake cultures ofCladosporium cladosporiodes, Aspergillus repens, A. ruber andPenicillium cyclopium as well as spores ofP. cyclopium andA. repens. The mites always produced fewer eggs on the fungal diets than on the ideal diet.Tyrophagus longior usually did best of the three mite species on the fungal diets, andG. destructor worst.Aspergillus ruber proved the most suitable fungus for all mite species, whileC. cladosporiodes was the least suitable. Spores were a less suitable diet than mycelial pellets from shake cultures, which were predominantly mycelium.     

Invertebrate biodiversity affects predator fitness and hence potential to control pests in crops

Natural enemies that control pests usually allow farmers to avoid, or reduce, the use of pesticides. However, modern farming practices, that maximize yields, are resulting in loss of biodiversity, particularly prey diversity. Does this matter? Pests continue to thrive, and without alternative prey the predators should, perforce, concentrate their attentions upon the pests.We showed that a diverse diet significantly enhances predator fecundity and survival. Experiments were conducted using common generalist predators found in arable fields in Europe, the carabid beetle Pterostichus melanarius (Coleoptera: Carabidae) and the linyphiid spider Erigone atra (Araneae: Linyphiidae). We tested the hypothesis that mixed species diets were optimal, compared with restricted diets, with respect to parameters such as predator weights, egg weights, numbers of eggs laid, egg development times, egg hatching rates and predator survival. In carabids, an exclusive earthworm diet was as good as mixed diets containing earthworms for egg production and hatching, but less good than such mixed diets for increase in beetle mass and sustained egg laying. For spiders, aphids alone (Sitobion avenae) or with the Collembola Folsomia candida, drastically reduced survival. Aphids plus the Collembola Isotoma anglicana improved survival but only aphids with a mixed Collembola diet maximized numbers of hatching eggs.Predators offered only pests (slugs or aphids) had lowest growth rates and fecundity. We therefore demonstrated that conservation of a diversity of prey species within farmland, allowing predators to exploit a diverse diet, is essential if predators are to continue to thrive in crops and regulate agricultural pests.     

Reproductive output of the ornamental shrimp Lysmata vittata (Stimpson, 1860) (Decapoda: Caridea) in wild populations and under different maturation diets

ABSTRACT

Reproductive output parameters (fecundity and egg volume) of Lysmata vittata were examined in a population in northeastern Brazil. Effect of maturation diets on the reproductive output of these shrimp under laboratory conditions was evaluated. Reproductive output was estimated for 25 shrimp collected in the wild. Another 45 pairs of shrimp were used for diet experiments, 15 pairs per treatment (T1: industrialized food, T2: fresh food, T3: mixed diets). For wild population, mean fecundity and egg volume were differed between developmental stages of the eggs. Fecundity was significantly lower in the specimens subjected to T1 diet (267 ± 141 eggs) compared with shrimp from the wild population (393 ± 183 eggs). Egg volume was significantly lower in shrimp subjected to three diets tested compared with those from wild population. Egg loss during embryonic development in L. vittata may be caused by several factors (e.g. aborted development and maternal cannibalism). Fresh food proved to be important for improving reproductive output in L. vittata reared in culture. We emphasize the significance of improving nutritional value and palatability of diets to improve cultivation efficiency. Lysmata vittata can be used as a model organism for future studies aiming to improve the cultivation techniques for shrimp of the genus.

Abbreviations PSH: protandric simultaneous hermaphroditism; ARS: artificial refuge structures; CL: carapace length; EV: egg volume; L: largest diameter of the egg; S: smallest diameter of the egg; T1: industrialized pellets; T2: fresh ingredients; T3: mixed diets; ANCOVA: analysis of covariancr; SD: standard deviation.     

Mollusc herbivory influenced by endophytic clavicipitaceous fungal infections in grasses

Clavicipitaceous fungi of the genus Neotyphodium occur widely as mutualistic, systemic, seed‐borne infections in festucoid grasses. Grass infection by these fungi is associated with the presence of a range of secondary metabolites (SM), several of which have been demonstrated to confer to the plant resistance against herbivorous vertebrates and insects. An initial experiment demonstrated that endophytic infection by Neotyphodium can influence the utilisation of grasses by Deroceras, with feeding preferences and impact on plant yields affected differentially by endophytes with different SM profiles. The role of Neotyphodium SM in feeding preferences of Deroceras slugs were then evaluated in artificial diets. Among the indole diterpenoids tested, lolitrem B was demonstrated to reduce feeding, while diets containing paxilline, lolitriol, α‐paxitriol and β‐paxitriol tended to be preferred over that of untreated diet. The pyrrolopyrazine alkaloid peramine had no effect. Among the ergopeptine alkaloids tested in the diets, ergotamine and ergovaline were demonstrated to be phagostimulatory. These results with artificial diets were generally consistent with Deroceras reticulatum preferences among plants of known Neotyphodium endophyte strain and SM profile. Deroceras slugs obtained from sites containing contrasting frequencies of Neotyphodium‐infected grasses, exhibited differential responses to Neotyphodium SM incorporated into artificial diet. This study demonstrates that infection of grasses by different isolates of Neotyphodium endophytes differentially influence herbivory by molluscs, reflecting their SM profile. These results offer an explanation for variable acceptability of grasses to molluscs and their importance in the diet of molluscs in the field reported in previous studies in both natural and agricultural systems. Neotyphodium endophytes potentially offer novel approaches to management of mollusc pests in agricultural gramineous crops.     

Influence of fungi on growth and survival of Onychiurus armatus (Collembola) in a metal polluted soil

Summary The influence of food quantity and quality on growth and survival of Onychiurus armatus (Tullb.) in metal polluted environments has been investigated in laboratory experiments. The Collembola was reared on five species of fungi isolated from a metal polluted soil close to a brass mill in SE Sweden.Survival of O. armatus was improved when fungal biomass was continuously added in a polluted mor (1,300 ppm Zn and 200 ppm Cu), and when specimens were fed metal polluted fungi for 1, 3 and 7 days a week, only those that were starved had increased mortality. Allometric growth, on the other hand, was significantly reduced when Collembola was given surplus of metal polluted fungi, whereas growth losses caused by metals were offset by protein rich food. Hence, sufficient food quantities alone could overcome mortality losses but not growth retardation in a metal polluted environment.Feeding preference of O. armatus was not determined by the protein content of the fungi although this was beneficial for growth. Metals changed the relative palatability of fungal species, but one of the metal tolerant species, Paecilomyces farinosus, which was also protein rich, remained reasonably attractive for O. armatus also when it was metal polluted. The mechanisms by which growth and survival of O. armatus were promoted by a combination of protein and Zn/Cu rich fungi seemed to be crucial in understanding the fate of a population of this species in a metal polluted soil.     

Fitness consequences for copepods feeding on a red tide dinoflagellate: deciphering the effects of nutritional value, toxicity, and feeding behavior

Phytoplankton exhibit a diversity of morphologies, nutritional values, and potential chemical defenses that could affect the feeding and fitness of zooplankton consumers. However, how phytoplankton traits shape plant–herbivore interactions in the marine plankton is not as well understood as for terrestrial or marine macrophytes and their grazers. The occurrence of blooms of marine dinoflagellates such as Karenia brevis suggests that, for uncertain reasons, grazers are unable to capitalize on, or control, this phytoplankton growth—making these systems appealing for testing mechanisms of grazing deterrence. Using the sympatric copepod Acartia tonsa, we conducted a mixed diet feeding experiment to test whether K. brevis is beneficial, toxic, nutritionally inadequate, or behaviorally rejected as food relative to the palatable and nutritionally adequate phytoplankter Rhodomonas lens. On diets rich in K. brevis, copepods experienced decreased survivorship and decreased egg production per female, but the percentage of eggs that hatched was unaffected. Although copepods showed a 6–17% preference for R. lens over K. brevis on some mixed diets, overall high ingestion rates eliminated the possibility that reduced copepod fitness was caused by copepods avoiding K. brevis, leaving nutritional inadequacy and toxicity as remaining hypotheses. Because egg production was dependent on the amount of R. lens consumed regardless of the amount of K. brevis eaten, there was no evidence that fitness costs were caused by K. brevis toxicity. Copepods limited to K. brevis ate 480% as much as those fed only R. lens, suggesting that copepods attempted to compensate for low food quality with increased quantity ingested. Our results indicate that K. brevis is a poor food for A. tonsa, probably due to nutritional inadequacy rather than toxicity, which could affect bloom dynamics in the Gulf of Mexico where these species co-occur.     

Interactions between pinewood nematodes and the fungal community of pine trees

The pinewood nematode Bursaphelenchus xylophilus, is the pathogenic agent of pine wilt disease and a globally notable pine pest. Despite being a plant pathogen, B. xylophilus has a mycophagous phase during its life cycle. We assessed the capacity for polyphagy of mycetophagous pinewood nematodes, testing which of the common species of fungi in pine trees provide better food and higher population growth rates. B. xylophilus performed particularly well on airborne fungi, namely the endophytes Botrytis cinerea and Cladosporium herbarum, and the pathogens Sirococcus conigenus and Sphaeropsis sapinea. Surprisingly, growth performance was not as good on the blue stain species (Ophiostoma spp. and Leptographium spp.) which are considered natural associates of B. xylophilus in the wild. Most of the fungi nonetheless permitted positive population growth of B. xylophilus, which is polyphagous and capable of feeding on numerous fungal species with diverse ecological niches.     

The effect of diet and temperature on reproductive rate in Arcella vulgaris Ehrenberg (Sarcodina: Testacida)

Summary The acceptability of bacterial, algal and fungal food cources as manifested by reproductive rate was investigated in Arcella vulgaris at 5°C intervals between 10°C-25°C. Bacteria and diets containing bacteria produced the highest rates of reproduction, a fungal diet induced the lowest rate of reproduction. On all diets reproductive rate increased with temperature up to 20°C, thereafter levelling out. Generation times were long, ranging from 64 h to 95 h, depending on diet and temperature. Consumption experiments using bacteria as food indicated that energy consumption was independent of temperature.