Critical periods in the life cycle and the effects of a severe spate vary markedly between four species of elmid beetles in a small stream

1. The chief objectives were: (i) to describe quantitatively the life cycles of four species of Elmidae, Elmis aenea, Esolus parallelepipedus, Oulimnius tuberculatus and Limnius volkmari; (ii) to use life tables to identify critical periods for survival in the life cycle of each species; (iii) to evaluate the immediate and longer‐term effects of a severe spate on densities of the four species. Monthly samples were taken over 63 months at two contrasting sites in a small stream: one in a deep section with macrophytes abundant, and the other in a shallow stony section. 2. There were five larval instars for O. tuberculatus, seven for L. volkmari and six for the other two species. The life cycle of each species took 1 year from egg hatching (chiefly in June for E. aenea and O. tuberculatus, and July for the other species) to pupation in the stream bank and a further year before the adults in the stream matured and laid their eggs. Mature adults were present in most months, but were rare or absent in January and February and attained maximum densities in April for O. tuberculatus and May for the other species. 3. Laboratory experiments provided data on egg hatching and pupation periods and the number of eggs laid per female. Life tables compared maximum numbers per square metre for key life‐stages. Within each species, mortality rates between adjacent life‐stages were fairly constant among six cohorts and between sites, in spite of large differences in numbers. The only exception for all species was the high adult, but not larval, mortality during a severe spate. 4. Standardised life tables, starting with 1000 eggs, identified key life‐stages with the highest mortality, namely the early life‐stages for E. aenea (36% mortality), start of the overwintering period to pupation for O. tuberculatus (41%) and L. volkmari (51%), start of pupation to the maximum number of immature adults for E. parallelepipedus (41%) and between the maximum numbers of immature and mature adults for O. tuberculatus (41%). Therefore, critical periods for survival in the life cycle differed between species, presumably because of their different ecological requirements. Similarly, the effects of the spate on adult mortality, and hence egg production, varied between species, being most severe and long‐term for E. aenea and O. tuberculatus, less severe for E. parallelepipedus and least severe with a rapid recovery for L. volkmari. Possible reasons for these discrepancies are discussed, but more data are required on the food and microhabitat requirements of the elmids before satisfactory explanations can be found.     

Mayfly production in a Colorado mountain stream: an assessment of methods for synchronous and non-synchronous species

Year-round collections of mayflies (Ephemeroptera) from a Colorado mountain stream allowed critical examination of several methods of calculating production for species with different life cycles. Five of the six numerically dominant species exhibited slow seasonal, univoltine life cycles. Baetis tricaudatus was bivoltine. Two species demonstrated well synchronized development, three species were poorly synchronized and a sixth was intermediate. Mean density and biomass data from each sampling date were used to ascertain the goodness-of-fit of each species to the Allen curve. It is proposed that such information can provide quantitative criteria for identifying species with well synchronized development and thereby determine when it is appropriate to directly apply cohort methods while avoiding time intensive body size (e.g. head width) measurements necessary for size-frequency analyses. In addition, these data demonstrate that species specific production varies with gross changes in elevation.     

Mortality of the terrestrial adult and aquatic nymphal life stages of Baetis vernus and Baetis rhodani in the Breitenbach, Germany (Insecta: Ephemeroptera)

1. The mortality of Baetis vernus Curtis and Baetis rhodani Pictet during the terrestrial-aerial and aquatic life stages was studied at the Breitenbach near Schlitz, Hesse, Germany. The number of females emerging from the stream was recorded with emergence traps. To estimate mortality of females of both species during terrestrial life, numbers of emerging females were compared with numbers of females returning to the stream for oviposition, as shown by numbers of egg masses found in the stream. 2. Mortality of female B. vernus during their terrestrial life stage was 98.8%. It was 91.2 and 96.6%, respectively, during the first and second generations of B. rhodani. 3. To estimate the mortality of both sexes of B. rhodani during the aquatic life stage, the number of eggs laid by the first generation was compared with the number of adults emerging during the second generation. 4. Mortality during the aquatic phase (both sexes combined) of B. rhodani was 91.1%. 5. Mortality during the terrestrial life stages was at least as high, if not higher than during the aquatic stages. Evidently, there is a considerable export of organic material into the terrestrial environment around the stream. Mortality during terrestrial life may be an important regulator of population size.     

Ecophysiological attributes of adult overwintering in insects: insights from a field study of the nut weevil,Curculio nucum

Abstract Diapausing insect species have evolved a great diversity of life cycles, although overwintering occurs at a single development stage within most species. Understanding why diapause has evolved towards a given life stage requires investigation of both the ecological and physiological attributes. Notably, it is suggested that adult overwintering is more energy‐demanding than larval overwintering but it brings fitness gains by allowing adults to be synchronized with their seasonal requisites through an early spring emergence. This hypothesis is tested in field conditions in the nut weevil Curculio nucum, whose life cycle comprises an obligate 2‐year, nonfeeding underground phase, including a larval, followed by an adult, overwintering. In this species, adult wintering leads to an early spring emergence; at first glance, however, this does not enhance synchronization between weevils and their host because adults emerge more than 1 month before starting to breed. It is suggested that adult overwintering ultimately evolved in response to the phenology of the host, by allowing females to oviposit in nuts before their full sclerotinization. Adult overwintering appears to be costly because adults postpone reproduction for 1 year, incur a significant weight loss and require feeding before egg laying. Surprisingly, lipids are unaffected during diapause, lipogenesis even being likely in the summer metamorphosis. These results suggest that the lipids involved in egg production may entirely come from the larval stages, whereas the other nutrients are acquired through adult feeding but this remains to be tested.     

Longitudinal zonation and life cycles of macrozoobenthos in the Mauerbach near Vienna,Austria

A total of 12277 benthic invertebrates were caught from November 1990 to October 1991 at the Mauerbach, a first to fourth order forest brook near Vienna, Austria, using a Surber sampler and a hand net. Five taxa comprised 96.6% of the catch: Diptera larvae (45.6%), Amphipoda (37.6%) and larvae of Ephemeroptera (6.4%), Plecoptera (5.3%) and Trichoptera (1.7%). Mean values of macrozoobenthos density ranged from 39514 specimens m⁻² at upstream sampling site 5 in December to only 286 specimens m⁻² at sampling site 3 in May, shortly after a severe flood. Based on benthic invertebrate population structure, sites 5 and 6 (situated near the source) were clearly separated from the downstream sites 1 to 4. At upstream sites, shredders comprised up to 71.7% of the total, whereas at downstream sites collectors were most abundant. In addition, the proportion of eucrenal species decreased from 12% at site 6 near the source to only 2% at site 1 near the mouth. Among the insect species studied in detail, most were univoltine except Ephemera danica (Ephemeroptera) and Sericostoma personatum (Trichoptera), which had a two-year life cycle.     

Generational shape shifting: changes in egg shape and size between sexual and asexual generations of a cyclically parthenogenic gall former

Successive generations of multivoltine species experience selection specific to the spatiotemporal environments encountered that may lead to adaptive divergence in reproductive traits among generations. To compare reproductive effort within and between generations, appropriate volumetric models, selected on the basis of the analysis of egg shape, are required to estimate the sizes (volumes) of individual eggs. We assessed the shape and estimated the volume of individual eggs produced by the temporally and spatially segregated sexual and asexual generations of the gall former, Belonocnema treatae Mayr (Hymenoptera: Cynipini: Cynipidae). Egg shape, indexed as the difference between the polar and equatorial axes of the ellipsoidal eggs, was independent of egg size, but differed between generations. The relationship of egg shape and female body size within and between generations confirmed that egg shape is an intrinsic property of each generation. Generational differences in egg shape then informed the selection of volumetric models to estimate egg size. We modeled asexual generation eggs as both spheres and prolate spheroids, and sexual generation eggs as both cylinders and prolate spheroids. Choice of volumetric model changed estimates of egg size within the asexual generation by 23% and within the sexual generation by 50%. Comparisons between generations based on the above models produced estimated differences in egg volume that ranged from 16 to 114%. In both generations, a prolate spheroid was the most parsimonious model of egg volume. Based on this model, sexual generation eggs averaged 43% larger than asexual generation eggs. The increased size of sexual eggs was achieved via conservation of the egg’s equatorial axis and elongation of the polar axis. The shift in egg shape between sexual and asexual B. treatae is the first documented dimorphism in an egg characteristic expressed between generations of a cyclically parthenogenic organism.     

Factors favouring the development and maintenance of outbreaks in an invasive leaf miner Cameraria ohridella (Lepidoptera: Gracillariidae): a life table study

1 Life tables were constructed for seven generations of the invasive horse‐chestnut leaf miner Cameraria ohridella in Switzerland, to assess the factors allowing the moth to build and maintain outbreak densities and to identify ‘gaps’ among their mortality factors that could be targeted by new control methods. The fecundity of females was calculated and the mortality factors affecting all developmental stages determined. 2 Females contained 106.6 eggs. In captivity, the mean number of eggs laid per female varied between 34 and 82 eggs, with a maximum of 184 eggs. Egg mortality was 18–75% and was mostly due to unknown factors. 3 The main mortality factors affecting larvae and pupae in the leaves were predation by birds and arthropods in all generation and leaf senescence in the autumn generation. Parasitoids killed less than 5% of the population. 4 Nearly 90% of overwintering pupae died in dead leaves, the majority of them being killed by earthworms and other leaf decomposers. As a result, the overwintering generations were the only ones showing a net rate of increase less than one. Non‐overwintering generations had net rates of increase between four and 11, allowing populations to build up from spring to autumn. 5 The persistence of high outbreak densities in Europe is explained by a combination of at least three factors: (i) high fecundity; (ii) multivoltinism; (iii) the unusually low impact of traditional natural enemies of leaf miners and particularly parasitoids. 6 Yearly variations in population densities could be at least partly explained by differences in larval mortality due to leaf senescence and intraspecific competition in the last generation in autumn.     

Thiamin dynamics during the adult life cycle of Atlantic salmon (Salmo salar)

Thiamin is an essential water-soluble B vitamin known for its wide range of metabolic functions and antioxidant properties. Over the past decades, reproductive failures induced by thiamin deficiency have been observed in several salmonid species worldwide, but it is unclear why this micronutrient deficiency arises. Few studies have compared thiamin concentrations in systems of salmonid populations with or without documented thiamin deficiency. Moreover, it is not well known whether and how thiamin concentration changes during the marine feeding phase and the spawning migration. Therefore, samples of Atlantic salmon (Salmo salar) were collected when actively feeding in the open Baltic Sea, after the sea migration to natal rivers, after river migration, and during the spawning period. To compare populations of Baltic salmon with systems without documented thiamin deficiency, a population of landlocked salmon located in Lake Vänern (Sweden) was sampled as well as salmon from Norwegian rivers draining into the North Atlantic Ocean. Results showed the highest mean thiamin concentrations in Lake Vänern salmon, followed by North Atlantic, and the lowest in Baltic populations. Therefore, salmon in the Baltic Sea seem to be consistently more constrained by thiamin than those in other systems. Condition factor and body length had little to no effect on thiamin concentrations in all systems, suggesting that there is no relation between the body condition of salmon and thiamin deficiency. In our large spatiotemporal comparison of salmon populations, thiamin concentrations declined toward spawning in all studied systems, suggesting that the reduction in thiamin concentration arises as a natural consequence of starvation rather than to be related to thiamin deficiency in the system. These results suggest that factors affecting accumulation during the marine feeding phase are key for understanding the thiamin deficiency in salmonids.     

Genetic variation and the performance of a mass‐reared parasitoid,Trichogramma pretiosum (Hymenoptera: Trichogrammatidae), in laboratory trials

During mass rearing, adaptation of biological control agents to the rearing environment is a potential problem. Using the parasitoid wasp Trichogramma pretiosum, the performance of 26 highly inbred lines, five composite ‘populations’ (created from the inbred lines) and one insectary‐reared population was compared using fertility life tables. Of the composite populations, three were created with maximal and identical genetic variation as a mixture of all 26 inbred lines, but these were then reared for a different number of generations (2, 6 or 17) before their performance was measured. The remaining two composite populations were created based on the performance of the individual inbred lines: one was a combination of two inbred lines with a high intrinsic rate of natural increase (rm), ‘high rm’; and the other was a combination of two lines with a ‘low rm’. High and low rm populations were reared for two generations prior to testing. Parameters measured were fertility, longevity and sex ratio. We found no difference between the maximally variable population reared for two generations and the ‘high rm’ population (rm = 0.285 and 0.282, respectively). ‘Low rm’ was the population with the lower performance (rm = 0.255). Genetically variable population reared for two generations for 48 h produced significantly more offspring than the populations reared for 6 and 17 generations. Hybrid population derived from the high‐rm lines did significantly better than that derived from the low‐rm lines. Low‐performance populations become more male based than high performance at 48 h. The potential benefits to improve population's performance using inbred lines for mass rearing are discussed.     

Reproductive value in a complex life cycle: heat tolerance of the pitcher-plant mosquito, Wyeomyia smithii

Because mortality accumulates with age, Fisher proposed that the strength of selection acting on survival should increase from birth up to the age of first reproduction. Hamilton later theorized that the strength of selection acting on survival should not change from birth to age at first reproduction. As organisms in nature do not live in uniform environments but, rather, experience periodic stress, we hypothesized that resistance to environmental stress should increase (Fisher) or remain constant (Hamilton) from birth to age at first reproduction. Using the pitcher-plant mosquito, Wyeomyia smithii, we imposed heat stress by simulating the passage of a warm-weather front at different pre-adult and adult stages. Contrary to either Fisher or Hamilton, stress tolerance declined from embryos to larvae to pupae to adults. Consequently, reproductive value appears to have been of little consequence in the evolution of stage-specific tolerance of heat stress in W. smithii.     

南昌地区灰飞虱的生活史、繁殖和越冬生物学特性

为了探明灰飞虱Laodelphax striatellus (Fallén)在南昌地区的生物学特性, 本研究在实验室和自然条件下系统调查了该虫的年生活史, 温度对其生长发育、 繁殖、 性比及翅型分化的影响, 及其越冬生物学。结果表明, 灰飞虱在南昌一年发生4~7代。在18~32℃, 卵的发育历期随温度升高逐渐缩短。若虫的发育历期在18~28℃随温度升高逐渐缩短, 但若虫的发育历期在30和32℃时显著长于28℃的发育历期（P=0.000﹤0.05）。越冬个体的若虫期为143~187 d。卵和若虫的发育起点温度分别为10.17℃和7.51℃。在室外, 7月中旬高温下孵化的第4代若虫的发育历期也明显延长, 显示了高温诱导的夏季休眠现象。在18~28℃, 产卵前期随温度升高而逐渐缩短, 当温度上升到30℃时, 其产卵前期比26和28℃下有所延长。产卵期在22℃最长, 30℃下最短。成虫在20~24℃下的寿命最长。在28℃下, 平均每雌产若虫量最大。不论在室内还是在室外, 雌雄比均接近1∶1。在18~32℃, 羽化的成虫均以长翅型占绝对优势。在自然条件下, 越冬代和第6代羽化的成虫以短翅型占优势, 其他各代仍以长翅型占优势。在自然条件下, 9月中旬孵化的若虫就有少量个体滞育越冬, 10月中旬后孵化的若虫全部进入越冬。越冬若虫的龄期为1-5龄。本研究为该虫发生的预测及有效防控提供基础资料。     

Life cycle of three stonefly species (Plecoptera) from an Apenninic stream (Italy) with the description of the nymph of Nemoura hesperiae

The life cycles of Isoperla grammatica, Amphinemura sulcicollis and Nemoura hesperiae are studied in a Northern Italy stream, sited in the Apennines. The three species show a univoltine life cycle and their development coincides approximately in the same period of the year, although the one of I. grammatica is longer. Possible egg dormancy could exist in the A. sulcicollis and N. hesperiae life cycles, but this needs to be proved. The three taxa show a spring flight period in the study area. Growth is almost constant through the life cycle, except in A. sulcicollis in which an increase is observed at the end of the development. N. hesperiae exhibits a faster growth than the other species. In addition, in this paper, the nymph of N. hesperiae is described and designed for the first time.     

Evolution of complex life cycles of amphibians: bridging the gap between metapopulation dynamics and life history evolution

Current evolutionary models for amphibian life cycles reflect tradeoffs in size-specific growth and mortality rates between the aquatic and terrestrial stages. A limitation of these models is that they do not incorporate evolutionary phenomena that are associated with metapopulation structure. In this work I address components of the evolution of complex life cycles (CLCs) that are tied to the metapopulation dynamics of amphibians that use seasonal wetlands that vary in hydroperiod. In particular, I describe how selection for the minimum length of the larval period affects metapopulation viability and the selection/migration equilibrium. Selection to increase the minimum length of the larval period functionally reduces the number of viable breeding sites on the landscape, increases the average distance between neighboring sites, and increases the risk of metapopulation extinction. Within a metapopulation, asymmetric gene flow between populations that are adapted to different hydroperiods tends to swamp local selection for long larval periods at sites with long hydroperiods. The evolutionary stability of CLCs of many species with metapopulation structure may reflect the fact that extremely small metamorphs cannot survive on land, while lineages with long larval periods incur a high risk of metapopulation extinction. I encourage theorists to more carefully consider how life history traits and metapopulation viability are related for these and other taxa.     

Density reductions by predatory trout increase adult size and fecundity of surviving caddisfly larvae in a detritus-based stream food web

1. In some situations, individuals surviving in environments where predation is intense can grow faster because the benefits of release from intraspecific competition outweigh costs associated with anti-predator responses. Whether these 'thinning' effects of predation occur in detritus-based food webs where resource renewal occurs independently of consumption by consumers was studied. We investigated how effects of predatory brown trout ( Salmo trutta ) on the larvae of the detritivorous stream caddisfly, Zelandopsyche ingens , influenced the size and fecundity of the caddisfly adults.
2. Trout substantially reduced the abundance of Z. ingens larvae, but adult male and female Z. ingens were significantly larger in trout streams compared to fishless streams. Females in trout streams had 33% more eggs than fishless stream females, and egg sizes were not significantly different. In mesocosms, Z. ingens larvae in low density treatments reflecting trout stream abundances grew significantly faster than larvae in high density treatments that were characteristic of fishless stream abundances. Non-lethal trout presence did not influence case building behaviour, feeding rates or growth or Z. ingens larvae, indicating non-lethal effects of predators were negligible.
3. Increased adult size and fecundity associated with trout stream individuals were probably a result of predator thinning of larval density indirectly releasing surviving Z. ingens from intraspecific competition. Thus, predator thinning did influence interactions between larvae in this detritus-based food web as larval growth was strongly density-dependent. However, extrapolating the total number of eggs potentially produced indicates the increased fecundity of females in trout streams would not compensate for losses of larvae to trout predation.     

Responses of adult Hydrellia lagarosiphon to a revised diet: implications for life cycle studies and laboratory culturing techniques

Preservation of an insect culture under laboratory conditions is essential for its study. Numerous diets have been developed for entomophagous insects undergoing screening as biological control agents in attempts to improve the nutritional quality of food provided in laboratory settings. However, less emphasis has been given to developing a more effective laboratory diet for phytophagous insects with the adult life stage not dependant on the target species. The larvae of Hydrellia lagarosiphon Deeming (Diptera: Ephydridae) mine the leaves of Lagarosiphon major (Ridl.) Moss ex Wager (Hydrocharitaceae). This species is currently under consideration as a biological control agent of this aquatic invasive. Rearing techniques for the adult stage of other Hydrellia species have been developed but current diets are not ideal as they result in relatively low reproduction rates. We compared alternative nutritional regimes and quantified their impact on life history attributes of H. lagarosiphon. The diets included the previously developed yeast‐sugar diet, a newly developed insect‐derived diet, and a diet that combined the two. Total fecundity was significantly higher for females on an insect‐derived diet compared with the traditional carbohydrate diet and the net reproductive rate (R_o) was also higher. Population doubling time (T_d) was lower, decreasing by 30% compared to the traditional laboratory diet developed for Hydrellia species. Adult females fed the combination diet, including both insect and non‐insect foods, laid 30% fewer eggs than those reared on an insect diet alone. Consequently, insect‐derived nutritional regimes could improve culturing techniques significantly and if permission to release the agent is granted, this diet may benefit mass rearing efforts potentially saving time and reducing associated costs.     

Bimodal life cycle of the burying beetle Nicrophorus quadripunctatus in relation to its summer reproductive diapause

Abstract 1. Under natural conditions in Kyoto, Japan, the reproductive activities of Nicrophorus quadripunctatus Kraatz (Coleoptera: Silphidae) decreased in summer and the species showed a bimodal life cycle.
2. In the laboratory, most adult pairs raised at 20 °C under a LD 12:12 h regime reproduced when provided with a piece of chicken. In adults raised at 20 °C under a LD 16:8 h regime, however, both reproductive behaviour and ovarian development were reduced. It is concluded that these adults entered a reproductive summer diapause.
3. High temperature (25 °C) also suppressed the reproductive behaviour even under a favourable LD 12:12 h regime. In the field, therefore, adults reduce their reproductive activity in summer because of diapause induced by long-day photoperiods and direct inhibition of reproduction by high temperatures.
4. When the temperature was changed from 20 °C to 25 °C immediately after hatching of larvae, they reached the wandering stage in 95% of adult pairs. When the temperature was changed from 20 °C to 25 °C immediately after oviposition, however, no larvae hatched in 85% of pairs. Egg mortality was significantly higher at 25 °C than at 20 and 22.5 °C; no eggs hatched at 27.5 °C. The physiological mechanisms for reducing reproduction probably prevent the beetles from inefficient oviposition in summer.     

Life cycle phenology of common detritivores from a temperate rainforest stream

The timing of life cycles, including growth rates, was determined for eight common species of detritivorous insects in a second-order stream in southwestern British Columbia, Canada. Six of the species (Zapada cinctipes, Z. haysi, Malenka californica, M. cornuta, Capnia sp., and Lepidostoma roafi) had simple, univoltine life cycles. The leuctrid stonefly Despaxia augusta has a 2-year life cycle, with an apparent egg diapause of about 6 months. The chironomid Brillia retifinis produced at least three generations per year. The major growth periods for the set of species considered here span the entire year. Adults of several species exhibited seasonal declines in size at emergence, but one species had larger adults as the emergence period proceeded. Closely related taxa had more similar life cycle timing than more distantly related species suggesting a degree of phylogenetic constraint in phenology of their life cycles. The influence of the timing of leaf drop on timing of life cycles for these animals does not fit with proposed scenarios based on fast and slow leaf processing rates.     

Some observations on the distribution and life history of Caenis robusta Eaton (Ephemeroptera) in Cheshire and North Shropshire,England

Caenis robusta Etn. was found in only 8 of 107 water bodies sampled. All the habitats from which it was recorded contained plant debris; the waters were still or slow-flowing and had conductivities ranging from 120–687 µmhos cm^–1 at 25°C.In North Shropshire, England, C. robusta had one generation per year. Newly hatched nymphs were abundant in August and growth occurred rapidly until September. No growth took place from October until March but was again rapid from April until June. Emergence took place at dusk in late June and July.