Life history and production of Caenis luctuosa (Burmeister) (Ephemeroptera, Caenidae) in two nearby reaches along a small stream

Population dynamics and production of C. luctuosa were compared in two reaches of the Agüera stream (northern Spain). This species showed univoltine winter life history in both sites. However, the start of the recruitment period, and the cohort production interval differed in 1 month between reaches. Secondary production of C. luctuosa ranged from 76 mg m^–2 year^–1 (upper site) to 93 mg m^–2 year^–1 (lower site). Although annual production seemed to be mainly influenced by the biomass found at each site, changes in life history may have also been important. The need to have accurate information about life history of the analysed species at the study sites when assessing secondary production is highlighted.     

Life history and secondary production of Electrogena ujhelyii (Ephemeroptera: Heptageniidae) in a small forest stream in West Carpathians

In this study we focused on life history parameters of Electrogena ujhelyii. We studied life cycle, nymphal growth pattern, biomass and secondary production in a small, calcareous stream in Malé Karpaty Mts (West Carpathians). The life cycle was univoltine with a large range in the size of nymphs during most samples dates. Mean annual density was 40 ind. m^?2 and the annual secondary production reached 183 mg DW m^?2 y^?1. There are no published data available on the secondary production values of this species, therefore these are the first published data.     

Life cycle and secondary production of two species of Campsurus (Ephemeroptera,Polymitarcyidae) in reservoirs of southeastern Brazil

The aim of this paper was to study the life cycle, the annual secondary production, and the spatial and temporal variation of the species Campsurus truncatus and C. violaceus in two reservoirs in Minas Gerais, Brazil. Campsurus truncatus was recorded in 11 months and had a multivoltine lifecycle with a secondary production of 4.61 g.m^?2.year^?1. Campsurus violaceus was collected in eight months and had a bivoltine lifecycle with a secondary productivity of 1.96 g.m^?2.year^?1.     

Life cycle and production of Skwala parallela (Frison) (Plecoptera: Perlodidae) in a Colorado montane stream

The life cycle and production of Skwala parallela, a perlodid stonefly, was investigated in a third-order Colorado montane stream. The species exhibited a univoltine life cycle with a distinct cohort. Small nymphs appeared in May. Rapid growth was exhibited throughout summer and autumn. During winter, growth slowed somewhat but was continuous until April. Maximum density of 34 nymphs/m² occurred in July. Based upon the instantaneous growth method, annual production was 395.3 mg/m² or 3.95 kg/ha dry weight with a P/B ratio of 4.4     

Studies on Ephemerella ignita (Poda) in a chalk stream in Southern England

The nymphs ofEphemerella ignita occurred in all months except October in the Bere Stream at Bere Heath, a small chalk stream in Southern England. Peak densities of 1,498 m^–2 and 753 m^–2 were recorded in May by two sampling methods used simultaneously. The seasonal patterns of growth and abundance are compared with the findings of other authors. A maximum biomass of 0.82 g m^–2 preserved dry weight is compared with figures for two of the most common invertebrate groups present; the Simuliidae and Oligochaeta.     

Life cycle ofPseudodiamesa branickii (Chironomidae) in a small upland stream

The generation time ofP. branickii was studied using larval samples in conjunction with rearing experiments and continuous collection of egg masses across one year. This species produced three generations per year in a central German stream (280 m a.s.l., 50° 40 N). Its generation time was variable and obviously influenced by the photoperiod to which eggs and larvulae were subjected. It is thus concluded that two strains ofP. branickii were present in a single population, one bivoltine and the other trivoltine.     

Migratory life cycle of Ephemerella maculata (Traver, 1934) (Ephemerellidae)

In this paper, I describe for the first time the migratory life cycle of an univoltine mayfly, Ephemerella maculata (Traver, 1934 Traver, J.R. (1934), ‘New North American Species of Mayflies (Ephemerida)’, Journal of the Elisha Mitchell Scientific Society, 50(1/2), 189–254. [Google Scholar]), between mainstems and tributaries of stream networks in California. Ephemerella maculata nymphs grow in sunny open mainstems and then emerge. Female adults fly into adjacent shaded tributaries, oviposit eggs, and die. The eggs diapause over summer and hatch in the fall in tributaries; then small nymphs drift down back to the mainstem before the subsequent spring growing season. Isotope analyses validate the adult migrations from mainstem to tributaries.

?Field rearing experiments show that the abundant algae in sunny mainstems are important food for the growth of E. maculata nymphs. However, eggs survive and hatch equally well in mainstem and tributaries, therefore, the fitness advantage of their migration to tributaries remains unclear.     

The distribution,life cycle and production of Leptophlebia vespertina (L.) (Ephemeroptera) in a lowland lake

Oak Mere, the water body studied, is a moderately productive base-poor lowland lake. It has zones of submerged marginal vegetation growing on sand at the south-west end and on peat at the north-east end. The distributions of L. vespertina and other macroinvertebrates are described in relation to these two distinct areas. Laboratory experiments on behaviour and survival are described in an attempt to explain the patterns of distribution. The seasonal distribution, variations in population density, spatial distribution variance in relation to mean density, life cycle, length-weight relationship, growth and biomass of L. vespertina are then described during two years at the sandy south-west end. The data are used to estimate annual production. Distribution, life cycle, growth and production are compared with populations in water bodies of lower conductivity in the British Isles and Scandinavia.     

Movement and mortality of stocked brown trout in a stream

The movement and mortality of stocked brown trout Salmo trutta were investigated using radio telemetry. Four brown trout left the study area whereas the remaining fish were stationary. After 5 weeks, 13 out of 50 tagged brown trout were still alive in the stream. Surviving fish had a significantly lower mean movement per day than fish, which later either died or disappeared. This difference in behaviour was most pronounced 2 to 8 days after release. Predation by the otter Lutra lutra was probably the main cause of the observed mortality.     

Life history of a lowland burrowing mayfly, Ephemera orientalis (Ephemeroptera: Ephemeridae), in a Korean stream

Life history aspects of Ephemera orientalis, a common lowland burrowing mayfly that resides in temperate East Asia, were studied in terms of voltinism, secondary production, and accumulated degree days for larval development. From March 1998 to June 1999, larvae were sampled monthly (weekly or biweekly during the emergence period) from a lower reach of the Gapyeong stream in Korea, a stream typical of temperate East Asia, using a Surber sampler (0.25 m², mesh 0.25 mm). As a result, the mean density of E. orientalis was 47.21 ± 13.58 indiv. m⁻² during the study period. Very small larvae less than 2 mm in body length were sampled on three separate occasions, and emergence was observed between late April and early October, except during late May. Based on the larval body length distribution and emergence time, two different developmental groups could be distinguished: the slow developmental group (S-group) and the fast developmental group (F-group). The F-group completed its life cycle within 4 months, whereas the S-group had a one-year life cycle. The S-group could be divided into two subgroups, dubbed the S1 and S2-groups, based on larval development. The developmental groups alternate their life cycles in the order: S1→F→S2→S1. Estimated annual production of the larvae was 68.81 mg DW m⁻² y⁻¹; mean biomass was 8.43 mg DW m⁻²; the annual production to mean biomass ratio was 8.16. The annual mean water temperature of the study year was 14.76 ± 6.63°C. The total accumulated degree days for larval development was 1396°C for the F-group, 2,055°C for the S1-group, and 1,975°C for the S2-group. Two different adult body size groups were distinguished (P < 0.001 by t-test): larger adults, which belonged to the S-group, were present throughout the emergence period, whereas smaller adults, which belonged to the F-group, were mainly present later in the emergence period (August–October). The difference in the accumulated degree days between the developmental groups may explain the size differences of adults.     

Life cycle, production, microdistribution and diet of the damselfly Euphaea decorata (Odonata: Euphaeidae) in a Hong Kong forest stream

Euphaea decorata in Tai Po Kau Forest Stream (Hong Kong) was univoltine. Most recruitment took place in summer, and larval growth proceeded throughout the year. Life-cycles recorded in 1977–78 and 1978–79 were similar. Annual production estimates, using the removal-summation, instantaneous growth and size-frequency methods, were more similar for the 1978–79 generation (ranging from 158.7–174.7; mean 1671 mg dry wt m^-2) than for the 1977–78 generation (93.9-173-6; mean 131.7 mg dry wt m^-2). Mean biomass was similar for both generations (ranging from 33.5–33.9 mg dry wt m^-2), and mean P/B ratios were 3.9 1977–78) and 5.0 (1978–79). These are the first estimates of annual production by an Oriental stream insect.
Larvae were most abundant at microsites in the middle of the stream. Multiple regression analysis indicated that substratum characteristics were a major determinant of microdistribution. Euphaea decorata apparently favoured poorly-sorted sediments with highly peaked grain size-frequency distributions, containing few fine particles.
The carnivorous larvae showed ontogenetic changes in diet. Small individuals consumed mainly chironomid (Diptera) larvae; the diet expanded to include (successively) larvae of Ephemeroptera and Trichoptera as E. decorata grew. Seasonal changes in diet were also apparent, although larval diets during spring and summer were similar. There was also considerable overlap between autumn and winter diets. Ontogenetic influences upon prey consumed were not sufficient to account for the observed seasonal differences.     

Seasonal variations in the benthos of a chalk trout stream,the River Samson,Belgium

Seasonal variations in the benthos of a Belgian chalk stream were investigated. Total biomass varies extensively over the year (30 to 100 g wet wt. m^–2). In summer, however, it is smaller than fish biomass. Densities range from 30 000 to 50 000 individuals m^–2, but the actual number of taxa is always near 60. Diversity indices (Shannon-Weaver and Pielou) exhibit slight fluctuations. Numerical indices are minimal in winter when the stream is in flood. Gravimetric indices are minimal in spring when biomass is maximal. Species-abundance curves almost always indicate a fauna with many rare species, with, also, some indication of a second maximum of moderately abundant species. Six taxa dominate: Oligochaeta, Gammarus fossarum, Ephemeroptera, Coleoptera, Trichoptera and Diptera. There are great fluctuations in their relative importance over a year. Thus, there seems to be a succession of peaks in densities of individual taxa permitting more efficient utilization of resources. However, biomass variations depend on the species involved. As a general rule, maximum biomass precedes maximum density.     

Habitat suitability modelling for mayflies (Ephemeroptera) in Flanders (Belgium)

Logistic regressions, artificial neural networks, support vector machines, random forests and classification trees were used to predict habitat suitability for mayflies on basis of physical–chemical water characteristics, dominant land use type and structural parameters. As a case-study, the dataset of Flanders was used, which consisted of 4289 samples containing mayflies (presences) and 3315 samples from waters where mayflies were never encountered (absences). For all techniques, data were randomly divided in a training set (two thirds) and a test set (one third). Models were calibrated using a tenfold cross-validation on the training set and subsequently validated using the test set. All techniques delivered good models that were able to discriminate sites with and without mayflies and performance (expressed as percent correctly classified instances and kappa-statistics) was in all cases similar for the training and the test set. Artificial neural networks and random forests performed slightly better compared to the other techniques. Samples with mayflies contained significantly more oxygen, a better developed river structure, higher values for sinuosity and steeper slopes, while samples without mayflies had significantly higher values for ammonium, nitrite, Kjeldahl nitrogen, total phosphorous, orthophosphate, biological and chemical oxygen demand, pH and conductivity. Also land use differed significantly, with mayflies usually present in forests but absent in industrial areas. The prevalence of mayflies gradually increased during the nineties from about 20 to 40%, which corresponded with an improvement of the chemical water quality. During the last decade, however, water quality did not further improve and as a result, mayflies prevalence did not continue to increase. Based on the planned measures, an ensemble forecast using the five mentioned modelling techniques predicted that mayflies prevalence will increase to 46% by 2015 and to 72% by 2027. To meet the requirements of the European Water Framework Directive, which states that all surface waters should obtain a good ecological quality, extra efforts will be needed to decrease nutrient concentrations and to improve habitat quality.     

The distribution of Ephemerella ignita (Ephemeroptera) in streams: the role of pH and food resources

SUMMARY. 1. The mayfly Ephemerella ignita does not occur in upland streams of the River Duddon catchment which have waters with low pH values of 4.8-5.2, negative alkalinities and low ionic contents. However, it does occur in lowland streams of the catchment which have waters with higher pH values of 6.6 and above, large positive alkalinities and high ionic contents. This study addresses the question of whether the absence of the animal in the acid water streams is due directly to the water chemistry or to the absence of suitable food.
2. In comparative experiments using unfed animals of several species it was shown that E. ignita has a considerable tolerance of low pH, low ion water. In this it is similar to Amphinemura sulcicollis (Plecoptera), a stonefly which occurs in the acid water streams of the River Duddon catchment.
3. Growth rates in laboratory experiments with E. ignita were equally good whether the food supplied was that available in the low pH water streams (bryophyte Nardia compressa plus filamentous alga Hormidium subtile) or typical of that available in high pH water streams (moss Rhynchostegium riparioides with the epiphytic diatom Cocconeis placentula). However, a consideration of the natural habits of the animal suggests that it is the inaccessibility of the H. subtile algal food, rather than its unsuitability, which is relevant to the exclusion of E. ignita from the low pH streams. In addition, the very limited diversity of other algae in the low pH water streams, in the summer season, may well be influential in inhibiting the establishment of E. ignita.
4. The comparative experiments using unfed animals suggested that, contrary to the findings for E. ignita , the mayflies Baetis muticus and Beatis rhodani are probably excluded from the acid water streams of the Upper Duddon because of water chemistry.     

Effect of temperature on the hatching time of eggs of Ephemerella ignita (Poda) (Ephemeroptera:Ephemerellidae)

SUMMARY. Eggs of Ephemerella ignita (Poda) were kept at eight constant temperatures (range 5.9–19.8°C) in the laboratory. Over 85% of the eggs hatched in the temperature range 10.0–14.2°C but the percentage decreased markedly to 39% at 5.9°C and 42% at 19.8°C. Hatching time (days after oviposition) decreased with increasing water temperature over the range 5.9–14.2°C and the relationship between the two variables was well described by a hyperbola. Therefore, the time taken for development was expressed in units of degree-days above a threshold temperature. Mean values (with 95%CL) were 552 (534–573) degree-days above 4.25°C for 10% of the eggs hatched, 862 (725–1064) degree-days above 3.57°C for 50% hatched and 1383 (1294–1486) degree-days above 3.14°C for 90% hatched. These values can be used to predict hatching times at temperatures below 14.68°C for 10% hatched, 14.54°C for 50% hatched and 14.45°C for 90% hatched. At higher temperatures, the hatching time and the number of degree-days required for development both increased with increasing temperature. Equations were developed to estimate the number of degree-days required for development at these higher temperatures.
Eggs were also placed in the Wilfin Beck, a small stony stream in the English Lake District. Maximum and minimum water temperatures were recorded in each week and the summation of degree-days was used to predict the dates on which 10%, 50% and 90% of the eggs should have hatched. There was good agreement between these estimates and the actual hatching times. Only 10–15% of the eggs hatched between October and late February with most of the eggs hatching in March, April and May. Nymphs hatching in October and November probably did not survive the winter.     

Seasonal dynamics of a mayfly (Insecta: Ephemeroptera) community in a Laurentian stream

The purpose of our study was to compare on a seasonal basis the emergence, the drift and the benthos on a short and uniform section (18.5 m)of a stream in the Laurentian Hills of Québec. Nineteen species occurred on this stretch, but only six were abundant: Leptophlebia cupida (Say), Habrophlebia vibrans Needham, Baetis pygmaeus (Hagen), Eurylophella verisimilis (McDunnough), Stenonema modestum (Banks), and Stenonema vicarium (Walker). Emergence was monitored for 5 years; densities differed from one trap to the next, as did species dominance, but total densities were similar. These data conform to Illies' rules of constancy and harmony. Nymphal densities were high (mean of 16000 m⁻²). The dominant species were univoltine, except H. vibrans (semivoltine), and B. pygmaeus (polyvoltine). Larger species contributed relatively more to the production of the reach despite their low densities than did the more abundant smaller species, especially those with long life cycles. Drift was low during most of the year, except during spring and species composition varied considerably. A comparison of the three data sets (emergence, drift and benthos) shows a general agreement for the less abundant species, but for three of the dominants, H. vibrans, L. cupida and E. verisimilis , movements of individuals to and from the reach at various times in the life cycle (adult flight, nymphal drift and migration) must be taken into account in order to correlate the data. None of the methods alone gives a satisfactory picture of the community structure, each is to some extent misleading and needs to be used in conjunction with the others; the conflicting results obtained from the data sets often reveal interesting life history traits which otherwise would not have been detected.     

Life history and production of Agapetus quadratus (Trichoptera: Glossosomatidae) in a temporary, spring-fed stream

1. The life history and trophic basis of production of the caddisfly grazer Agapetus quadratus were studied in the torrent Gorg Blau, a spring‐fed stream on the island of Majorca that dries annually during summer. 2. Quantitative random samples were taken every 2–3 weeks during an annual surficial flow period, from November 2000 to mid‐July 2001. Instars of field‐collected larvae were determined by measurements of head width and pronotum length, and the sex of all pupae was determined to study sexual dimorphism and sex ratio. 3. Stage‐frequency histograms suggested a trivoltine population, with an average cohort time of 4 months. Larval development was asynchronous, with continuous growth and overlapping generations. Recruitment peaks were identified in mid‐November, early March and late June, indicative of winter, spring and summer generations. On average, females were larger than males and the mean sex ratio was 2 : 3 (females : males). Population densities and biomasses derived from the field data were used to calculate production and turnover rate. 4. Annual production of A. quadratus in the torrent Gorg Blau (4.80 g dry mass m^?2 year^?1) was the highest ever reported for the genus, being comparable with that estimated for some insects with rapid development and multiple cohorts. 5. Estimates of production of A. quadratus were combined with foregut content analysis to estimate the fraction of total production derived from the principal food sources: algae and organic detritus. Algae supported a major proportion of the production of this grazer. 6. The low density of predators characteristic of many temporary streams, and the small amplitudes in discharge and temperature during most of the wet period that characterise the spring habitats might allow high levels of grazer production in this particular Mediterranean stream.     

Life histories, production dynamics and resource utilisation of mayflies (Ephemeroptera) in two tropical Asian forest streams

SUMMARY 1. A 2‐year study of the life histories, production dynamics and resource utilisation of five mayfly species was undertaken in two forest streams in Hong Kong [Tai Po Kau Forest Stream (TPKFS) and Shing Mun River (SMR)]. Afronurus sp. and Cinygmina sp. (Heptageniidae), Procloeon sp. and Baetiella pseudofrequenta (Baetidae), and Choroterpes sp. (Leptophlebiidae) were abundant in both streams and contributed more than 50% of the total mayfly populations. 2. All species had asynchronous larval development with recruitment occurring throughout the year. Mean annual production (all mayflies combined) was 3.1 and 2.0 g dry weight m^?2 year^?1 in SMR and TPKFS, respectively – the higher value at SMR reflecting greater mayfly densities – with more than 70% of production occurring during the wet season. Mayfly production varied between years, decreasing by 5% in TPKFS and 43% in SMR during 1996–97, reflecting lower densities of heptageniids relative to 1995–96. Annual biomass turnover rates (P/B) were high in both sites ranging from 27.2 to 94.6 in TPKFS (Cinygmina sp. and Procloeon sp.) and from 31.8 to 109.8 in SMR (Cinygmina sp. and B. pseudofrequenta). 3. Patterns of daily production in both streams showed that Afronurus sp., Cinygmina sp. and Choroterpes sp. were most productive during the wet season, while Procloeon sp. maintained high production levels throughout the year. The highest daily production of B. pseudofrequenta occurred during the wet season in TPKFS, but in the dry season at SMR. Temporal overlap in production and hence resource utilisation in both streams, calculated using the proportional similarity index (PS), ranged from 0.39 to 0.81. It was highest (0.63–0.81) between pairs of species of Heptageniidae and Baetidae, and lowest between Choroterpes sp. and other mayflies (0.39–0.61). No clear temporal segregation was observed among any species. However, when using the fraction of production attributable to each food, lower PS values were obtained for all species in both sites. In SMR, trophic segregation may have occurred between the two species pairs Procloeon sp.–Cinygmina sp. and Procloeon sp.–Choroterpes sp. (PS=0.17 and 0.03, respectively). 4. A combination of production data and information on the stable isotope signature of mayflies revealed that, during both the wet and dry seasons, more than 50% of total mayfly production in TPKFS was derived from autochthonous foods. In SMR, 68% of production was supported by allochthonous foods during the wet season, and 72% by autochthonous sources in the dry season. Considering that more than 70% of the total production occurred in the wet season, the trophic basis of mayfly production in SMR is mostly allochthonous (58%) while in TPKFS it is mainly of autochthonous origin (66%). The year‐round importance of autochthonous foods in shaded streams such as TPKFS is surprising, but the wet season contribution of allochthonous foods (especially in SMR) may have resulted from depletion of algal biomass during spates.