Instar sizes,life cycles,and food habits of five Rhyacophila (Trichoptera: Rhyacophilidae) species from the Appalachian Mountains of South Carolina,U.S.A.

The larval head widths at each instar, life cycles, and food habits of late instars were determined for five species of Rhyacophila from two Appalachian mountain streams in South Carolina, U.S.A. Rhyacophila acutiloba Morse & Ross was univoltine with two cohorts, one emerging in the spring and another presumably emerging in early autumn. Rhyacophila fuscula (Walker), R. nigrita Banks, and R. carolina Banks were apparently multicohort, univoltine species with extended flight periods. Rhyacophila minor Banks was univoltine with a spring emergence. All species were predaceous and consumed mainly Plecoptera nymphs and Trichoptera larvae.     

Typology of life cycles of ground beetles (Coleoptera,Carabidae) in Western Palaearctic

An original classification of the life cycles of ground beetles from Western Palaearctic is proposed. The classification is based on a combination of five criteria: duration, number of generations per season, phenology of reproduction, stability, and repeatability of reproduction. According to the individual lifespan, the cycles are subdivided into annual and biennial ones. The annual life cycles may be uni-and bivoltine, whereas biennial ones are always univoltine. By the time of reproduction, winter-spring, spring, spring-summer, early summer, summer, late summer, summer-autumnal, autumnal, autumn-winter, winter, and aseasonal species are distinguished. The biennial and bivoltine cycles may be of both facultative and obligate nature. Species living only one season and having a continuous reproductive period are designated as semelparous, while those breeding during two or more years or having several distinct periods of reproduction in one season, as iteroparous. By now, 30 variants of life cycles in Carabidae from western Palaearctic have been established. Repeated similarly directed modifications of the life cycle may produce essentially different seasonal rhythms in some individuals. In this case, two subpopulation groups usually appear within the population. Under the most unfavorable conditions, these groups become practically isolated and hibernate at different ontogenetic stages. The individual development in each of these groups takes two years with the same seasonal rhythm. Among the types considered, only obligate-bivoltine life cycles are always polyvariant, but annual univoltine and obligate-biennial ones are always univariant. The facultative-bivoltine and biennial life cycles may be realized as uni-and polyvariant ones, depending on the environmental conditions.     

Leaf age and season influence the relationships between leaf nitrogen, leaf mass per area and photosynthesis in maple and oak trees

Abstract. Seasonal changes in photosynthesis, leaf nitrogen (N) contents and leaf mass per area (LMA) were observed over three growing seasons in open-grown sun-lit leaves of red maple ( Acer rubrum ), sugar maple ( A. sacchamm ) and northern pin oak ( Quereus ellipsoidalis ) trees in southern Wisconsin. Net photosynthesis and leaf N were highly linearly correlated on both mass and area bases within all species from late spring until leaf senescence in fall. Very early in the growing season leaves had high N concentrations, but low photosynthetic rates per unit leaf N, suggesting that leaves were not fully functionally developed at that time. Leaf N per unit area and LMA had nonparallel seasonal patterns, resulting in differing relationships between leaf N/area and LMA in the "early versus late growing season. As a result of differences in seasonal patterns between leaf N/area and LMA, net photosynthesis/area was higher for a given LMA in the spring than fall, and the overall relationships between these two parameters were poor.     

Embryonic developmental rates of northern grasshoppers (Orthoptera: Acrididae): implications for climate change and habitat management

Accurate models of temperature-dependent embryonic developmental rates are important to assess the effects of a changing climate on insect life cycles and to suggest methods of population management by habitat manipulation. Embryonic development determines the life cycle of many species of grasshoppers, which, in cold climates, spend two winters in the egg stage. Increasing temperatures associated with climate change in the subarctic could potentiate a switch to a univoltine life cycle. However, egg hatch could be delayed by maintaining a closed vegetative canopy, which would lower soil temperatures by shading the soil surface. Prediapause and postdiapause embryonic developmental rates were measured in the laboratory over a wide range of temperatures for Melanoplus borealis Fieber and Melanoplus sanguinipes F. (Orthoptera: Acrididae) A model was fit to the data and used to predict dates of egg hatch in the spring and prediapause development in the fall under different temperature regimens. Actual soil temperatures were recorded at several locations over 5 yr. To simulate climate warming, 2, 3, or 4°C was added to each hourly recorded temperature. Results suggest that a 2, 3, or 4°C increase in soil temperatures will result in eggs hatching ≈ 3, 5, or 7 d earlier, respectively. An increase of 3°C would be required to advance prediapause development enough to allow for a portion of the population to be univoltine in warmer years. To simulate shading, 2 and 4°C were subtracted from observed temperatures. A 4°C decrease in temperatures could potentially delay hatch by 8 d.     

Dormant stages and their participation in adjustment and regulation of life cycles of harvestmen (Arachnida,Opiliones)

Information on seasonal adaptations in the life cycles of harvestmen (order Opiliones) summarized in this review reveals a great diversity of these arachnids with respect to duration, voltinism, and phenology of their life cycles, as well as to the number and ontogenetic position of the dormant stages required for survival during the winter season and also for synchronizing development with seasonal climate rhythms. Most harvestmen have stenochronous development with univoltine life cycles synchronized by arrest of development in hibernating eggs or (rarely) diapausing nymphs and adults. The number and arrangement of dormant stages represent stable species-specific traits, with some rare cases of interpopulation differences (e.g., in Phalangium opilio). Eurychronous harvestmen exhibit the year-round occurrence of main ontogenetic stages which show equal abilities for either active development or quiescence (depending upon the external factors). Two alternative types of development arrest are common in univoltine opilionids: (1) diapause induced at the early stage of embryogenesis and terminated during cooling (in Opilio parietinus and eleven other species) with transformation into postdiapause quiescence; (2) cold quiescence enforced by low temperatures at the last stage of egg development just before hatching (in Phalangium opilio and four other species). In conclusion, the systems of seasonal adaptations in Opiliones are compared with those in other arachnids, insects, and crustaceans. Some promising directions in the study of seasonal adaptations in opilionid life cycles are suggested.     

The life cycles of Leptophlebia vespertina (L.) and L. marginata (L.) (Ephemeroptera) in Llyn Dinas, North Wales

Data is presented on the life cycles of Leptopblehia vespertina (L.) and L. marginata (L.) in Llyn Dinas, an oligotrophic North Wales lake. Both species are univoltine and growth continues throughout the winter although somewhat retarded. Except during July and August there is a clear size separation of the two species, L. marginata being larger and emerging earlier. The egg incubation period was determined directly from laboratory studies. The results are discussed in relation to previous research in the English Lake District.     

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.     

Diet and predation by three leaf-associated stoneflies (Plecoptera) in an Arkansas mountain stream

SUMMARY. 1. Life cycles, food habits, mouthpart morphologies, prey preferences, and predator-prey size relationships were investigated for the three most common stoneflies in leaf packs in the Little Missouri River, Arkansas: Clioperla clio (Newman), Isoperla namata Prison, and Perlesta spp. Each species was univoltine and had a fast seasonal cycle, with extended egg or nymphal diapause, rapid nymphal growth and emergence in spring to early summer.
2. Foregut analysis showed predictable dietary shifts by C. clio and Perlesta spp., from detritus and diatoms in early instars to invertebrate prey in later instars. Isoperla namata fed facultatively on insects, detritus and diatoms, with no conspicuous ontogenetic shifts. The mouthparts of all three species strongly reflected their diet.
3. Electivity analysis indicated opportunistic feeding by the three species, with feeding rarely differing significantly from random.
4. Sizes of stonefly predators and their ingested prey were highly correlated ( P <0.01). However, rather than shifting toward larger prey sizes, growing predators expanded their size thresholds and continued to include numerous small prey items in their diet. Prey choice was governed most directly by prey availability, because stoneflies selected the most abundant groups, irrespective of size or taxon.
5. The likelihood of competition among the three stoneflies for prey was minimized by timing of their life cycles, differential food use and probably the temporary nature of leaf pack habitats.     

Fitness consequences of alternative life histories in water striders,Aquarius remigis (Heteroptera: Gerridae)

Using field and laboratory observations and experiments over 3 years, I investigated whether reproductive trade-offs shape individual life histories in two natural populations of the water strider, Aquarius remigis, in which univoltine and bivoltine life cycles coexist. Both later eclosion dates and food shortages, even after adult eclosion, induced diapause in females, thus deferring reproduction to the following spring. Adult body size was positively affected by food availability during juvenile development. Higher food levels also increased the reproductive output of females, but not their longevity or oviposition period. When compared to spring breeders (univoltine life cycle), direct (summer) breeders (bivoltine life cycle) experienced reduced lifetime egg numbers and longevity, as well as reduced survivorship of their second-summer-generation offspring; these reproductive costs offset, at least in part, the advantage in non-decreasing populations of having two generations per year. Fecundity was correlated with body size, and among summer-generation females direct breeders were larger than non-breeders. The time remaining before the onset of winter and/or the time since adult eclosion augmented cumulative energy uptake, and consequently the lipid reserves and winter survival probability of non-breeding (diapausing) summer adults approaching hibernation. Overwintered spring reproductives died at faster rates than non-reproductive summer individuals despite greater food availability in spring, indicating a mortality cost of reproduction. Body length correlated with absolute and not with proportional lipid content but showed no consistent relationship with survivorship in the field. These results are in agreement with current theory on the evolution of insect voltinism patterns, and further indicate high degrees of life history flexibility (phenotypic plasticity) in the study populations in response to variable environmental factors (notably photoperiod and food availability). This may be related to their location in a geographic transition zone from uni- to bivoltine life cycles.     

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.     

Species composition, host association and niche differentiation in fleas of small mammals in the Ilmen-Volkhov lowland

Species composition, abundance, annual cycles, and host association of fleas parasitizing small mammals were investigated. The problem of niche differentiation in these insects is considered on the base of the comparative analysis of their annual cycles. The annual cycles of the fleas are revealed to be similar in the case of few number of flea species in parasite community. Thus, two species parasitizing Sorex araneus (Doratopsylla dasycnema and Palaeopsylla soricis), as well as three species associated with Apodemus uralensis (Megabothris turbidus, Ctenophthalmus agyrtes, and Ct. uncinatus) have equal phenology of parasitizing. The fleas community of Clethrionomys glareolus is characterized by a large species number and high diversity of the annual cycles. The differentiation by the seasons of parasitizing is observed most clearly in the dominant flea species, namely Amalaraeus penicilliger, Ct. uncinatus, and Peromyscopsylla bidentata. The periods of imaginal life are overlapped significanly in these species, but they are differed by the season of dominance. Ct. uncinatus predominates in spring and summer, while P. bidentata predominates in autumn, and A. penicilliger predominates in winter and early spring. It may be noted also, that niche partitioning was not observed in the fleas having wide range of hosts. The imaginal life of such fleas usually does not go beyond the warm season.     

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.     

Description of a new Dendrocerus species (Hymenoptera: Megaspilidae) hyperparasitic on Stomaphis aphids and additional notes on their primary parasitoid, Protaphidius nawaii (Braconidae), and another hyperparasitoid, Euneura stomaphidis (Pteromalidae)

A food chain through three trophic levels on certain trees, Stomaphis aphids – a primary parasitoid, Protaphidius nawaii – a hyperparasitoid, Euneura stomaphidis , has been known in Japan. In the course of my survey on the seasonal prevalence of three Stomaphis species and their parasitoids from 1981–2007 in Kyoto, Japan, I found another hyperparasitoid, which I describe here as a new species, Dendrocerus stomaphis. From fragmentary observations, I inferred the life cycles of P. nawaii, E. stomaphidis and D. stomaphis . These three species have different strategies to escape ant aggression: P. nawaii by specific morphological and behavioral adaptation, E. stomaphidis by general anti-predator behavior like quick movement and D. stomaphis by a unique univoltine life cycle with the active phase from late autumn to early winter when the ants are less active.     

The role of litter in an old-field community: impact of litter quantity in different seasons on plant species richness and abundance

Summary We studied the effect of removing and adding plant litter in different seasons on biomass, density, and species richness in a Solidago dominated old-field community in New Jersey, USA. We removed all the naturally accumulated plant litter in November (658 g/m²) and in May (856 g/m²) and doubled the amount of litter in November and May in replicated plots (1 m²). An equal number of plots were left as controls. Litter removal and addition had little impact on total plant biomass or individual species biomass in the growing season following the manipulations. Litter removal, however, significantly increased plant densities but this varied depending upon the season of litter removal, species, and life history type. Specifically, the fall litter removal had a much greater impact than the spring litter removal suggesting that litter has its greatest impact after plant senescence in the fall and prior to major periods of early plant growth in spring. Annual species showed the greatest response, especially early in the growing season. Both spring and fall litter removal significantly increased species richness throughout the study. Litter additions in both spring and fall reduced both plant densities and species richness in June, but these differences disappeared near the end of the growing season in September. We concluded than in productive communities where litter accumulation may be substantial, litter may promote low species richness and plant density. This explanation does not invoke resource competition for the decline in species richness. Finally, we hypothesize that there may be broad thresholds of litter accumulation in different community types that may act to either increase or decrease plant yield and diversity.     

Comparison of the life cycles of two mayfly species between upper and lower parts of the same stream

The life cycles of the burrowing mayflies, Ephemera strigata Eaton and E. japonica McLach‐lan were studied comparatively between upper part and lower part of a small tributary of the Yoshino River. E. strigata was more common than E. japonica in the studies areas. E. japonica was found only in the upper part and in tiny tributaries of the stream. E. strigata was univoltine in the lower and in the upper parts of the stream. However, the population falls into two groups, one of which emerges in early May and the other in summer. Water temperature seems to be one of the factors affecting the variable life cycle of E. strigata. E. japonica is univoltine. Emergence of this species was observed from June to August.     

Distribution and life cycles of cladocerans in temporary pools from Central Italy

SUMMARY 1. A study of twenty-nine temporary pools located in a nature reserve in Central Italy was carried out for two successive years (1982–83 and 1983–84) to determine the distribution and life cycles of the cladocerans they contained.
2. The pools followed a typical autumnal cycle, filling up in early autumn and drying up in late spring. They differed in morphology (size, depth, type of soil) and in the length of the wet phase.
3. Nineteen cladoceran species were found. A few of these are early species, appearing at the beginning of the wet phase, whereas others appear much later in spring and these populations persist for a shorter time.
4. Species distributions did not show any clear correlation with the ponds'morphological features, and the life cycles of individual species were similar in all ponds. Thus populations did not adapt life history patterns to match conditions in particular ponds.     

Emergence patterns of univoltine and bivoltine Ips typographus (L.) populations and associated natural enemies

Control measures aiming at reducing bark beetle populations and preserving their natural enemies require a sound knowledge on their overwintering and emergence behaviour. These behavioural traits were investigated in univoltine and bivoltine populations of the European spruce bark beetle (Ips typographus [L.], Coleoptera: Scolytinae) and its predators and parasitoids over several consecutive years. In univoltine populations, roughly 50% of the bark beetles left their brood trees in fall together with most parasitoids and some significant predatory flies and beetles. In bivoltine populations, <10% of the second bark beetle generation emerged in fall and the remainder overwintered under the bark of their brood trees. Likewise, most predatory beetles and flies spent wintertime with their prey under the bark, while most parasitic wasps emerged in fall. The spring emergence of bivoltine predatory beetles was found to occur up to 3 weeks earlier than that of I. typographus, while that of the predatory flies and the parasitoids was delayed by up to 1 month. In univoltine populations, the bark beetles emerged several weeks prior to most antagonistic taxa. In the heat year 2003, three I. typographus generations were produced at the lower location, 36% of the third generation emerged in fall, while the proportions of overwintering predators remained largely the same as in previous years. Similar to their host, more parasitoids left their brood trees in fall after warm years. The results show that sanitation felling during winter probably kills most bark beetles in bivoltine populations, but also eliminates many natural enemies. In univoltine populations, sanitation felling might be less detrimental to both I. typographus and natural enemies because a fair fraction of their populations will already have left the trees before cutting. Warmer climates may affect the interactions of bark beetles and natural enemies and thus the impact of control measures.     

Seasonality and life cycles of woody plant-feeding noctuid moths (Lepidoptera: Noctuidae) in Mediterranean habitats

Abstract.

• 1 The annual cycle of noctuid moths feeding as larvae on the foliage of eighteen woody plant species (belonging to ten different families) was studied at two southeastern Spanish Mediterranean forest habitats. Two questions were addressed in this study. (1) Does the rigorous (hot and dry) summer season characteristic of the Mediterranean-type climate impose a constraint on noctuid life histories? (2) Are there detectable differences in abundance or phenological patterns between the noctuid assemblages feeding on evergreen and winter-deciduous trees and shrubs?
• 2 Regardless of their leaf persistence habit, the majority of woody species studied had short shoot growth and leaf production periods in spring. Abundance of noctuid larvae peaked in May-June, shortly after host plants started to produce new leaves. Virtually no noctuid larvae were found on the foliage of the woody species studied at other times of year. Abundance and seasonal pattern of occurrence of larvae were similar on evergreen and deciduous food plants.
• 3 In contrast with the unimodal seasonal pattern exhibited by larvae, the abundance of adult moths (assessed by light trapping) showed two distinct peaks in early summer and early autumn, and a marked minimum in mid-summer.
• 4 Most woody plant-feeding noctuids recorded in this study (86% of species, about 95% of individuals) were univoltine. Univoltine species fell into one of two distinct life history categories, corresponding to pre- and post-summer flight periods. These two contrasting phenologies were associated with a bimodality in the duration of the pupal stage (‘short’ versus ‘long’), and were closely related to taxonomical affiliation at the subfamily level.
• 5 Within the ‘short’ pupal duration group, some species mate and oviposit shortly after emergence and pass the dry season in the egg stage, while others remain as potentially active, non-reproductive adults over most of the summer and mate by the end of that season. Species in the ‘long’ pupal duration group pass the summer in pupal or prepupal stage.
• 6 In the Mediterranean habitats studied, the adversity of the summer dry season seems to have led to a woody plant-feeding noctuid species assemblage almost entirely made up of univoltine species that concentrate their larval phases when suitable food (young leaves) is most abundant, and ‘avoid’ mid-summer as a flight time by emerging either shortly before or shortly after the adverse summer drought period. Year-round foliage availability afforded by dominant evergreen plants does not seem to have influenced the seasonal organization of noctuid cycles in any substantial way.

     

Life history traits of Pseudasphondylia rokuharensis (Diptera: Cecidomyiidae) affecting emergence of adults and synchronization with host plant phenology

To identify adaptive strategies of gall-inducing cecidomyiids, we studied the life history traits of Pseudasphondylia rokuharensis (Diptera: Cecidomyiidae), a univoltine species inducing fruit galls on a deciduous shrub, Viburnum dilatatum. Although univoltine gall midges that are associated with trees or shrubs are usually difficult to rear from immature stages under artificial conditions, we successfully reared P. rokuharensis from first instar to adult. Mature larvae entered diapause in autumn, and the diapause terminated after low temperature treatment (4 mo of incubation at 5 degrees C). The adults emerged within a shorter period of time than those without low temperature treatment. Thus, the diapause provides better synchronization of adult emergence in the spring. Because adults live for only a few days, this synchronization improves the chances of adults finding a mate. This is the first report on the mechanism of larval diapause termination for univoltine and tree- or shrub-associated gall midges. The numbers of days needed for adults to emerge under laboratory conditions after low temperature incubation were significantly different between two different localities. This result might suggest the possible existence of ecoclines in their life history parameters.