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.     

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.     

The microdistribution of mayflies (Ephemeroptera) in myriophyllum beds in Pennington Creek,Johnston County,Oklahoma

I studied the population densities and the microdistributions of the four most abundant mayfly (Ephemeroptera) nymphs in Myriophyllum heterophyllum beds in Pennington Creek, Johnston County, Oklahoma, from May to December, 1977. The section of the stream studied has relatively constant flow rates and temperatures throughout the year due to continuous inflow from groundwater. This uniform physical environment allows the plant beds to grow throughout the year and provide a permanent habitat. The four mayflies are Tricorythodes fictus (Tricorythidae), Caenis delicata (Caenidae), Baetis favistriga and B. quilleri (Baetidae). All four species are of similar size and feed by scraping periphyton from the surface of the Myriophyllum. T. fictus and the two species of Baetis have overlapping microdistributions in the leafy, upper one-half of the plant bed. T. fictus and C. delicata have overlapping microdistributions in the lower portion of the plant bed and roots. Each species has similar population densities where their distributions overlap. There is no significant selection for position with respect to current flow in the Myriophyllum for any of the species. Food and space are always abundant, especially in the top one-half of the Myriophyllum. There are no major changes in these trends over the year.     

Life histories, microdistribution and drift of two mayfly (Ephemeroptera) species in the Pembina River, Alberta, Canada

Ephemerella inermis Eaton is a univoltine species that emerges in July. Eggs hatch in August and larvae grow rapidly in autumn and spring, but not during winter. Baetis tricaudatus Dodds produces three cohorts per year. Emergence periods occur in early June, July and early September. Only one cohort overwinters as larvae.
Microdistribution of both species is controlled largely by the discharge pattern of the river. During 1978, a year of frequent floods, animals were equally distributed among slow and fast water portions of the river. Nocturnal drift densities of both species were much greater in slow than in fast water areas. During 1979, a year of relatively stable flow, benthic larval densities were significantly greater in fast water than in slow water.
Early instar E. inermis larvae are most abundant in nearshore areas, but move to deeper water in late autumn. Baetis tricaudatus larvae are seldom numerous in near-shore areas.
Densities of both species were positively associated with concentrations of detritus and of other invertebrates, but only when relationships were considered in areas of similar current velocity independently of season.     

Life histories of burrowing mayflies (Hexagenia limbata and H. rigida, Ephemeroptera: Ephemeridae) in a northern Canadian reservoir

1. The life histories and population dynamics of the burrowing mayflies Hexagenia Umbata and H. rigida were examined from 1986 to 1988 in four regions of an impounded lake in northern Manitoba, Canada. 2. Bottom temperatures reached 15–20°C, but ice-free periods of 4–6 months resulted in annual degree day (dd > 10°C) accumulations of only 210–650 in various regions and years. 3. The life cycles (egg to adult) of both Hexagenia species required 3yr in the three warmest regions studied and 4yr in the coolest area; total dd accumulations to complete the life cycle ranged from 1222 to 1468 dd > 10°C. 4. Fecundity (no. of eggs/) was strongly correlated to body size of subimagos, and subimago size was strongly correlated to length of nymphal exuviae, so field collection of nymphal exuviae could be used to determine both the size and fecundity of the emerging females. 5. Degree day accumulation in the final year before emergence was a better predictor of emergence timing than overall dd accumulations for the life cycle.     

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.     

Life cycle, growth, mortality and production of Ephemerella major Klapalek (Ephemeroptera) in a trout stream in Belgium

SUMMARY. 1. The life cycle of Ephemerella major Klapalek in a chalk trout stream in Belgium took 1 year. Emergence was highly synchronized with a flight period from mid-May to mid-June. Tiny nymphs occurred from June to late August.
2. The mean instantaneous growth rate was high in autumn (3.6% wet wt day⁻¹), very low from November to February (0.8% wet wt day⁻¹) and high until emergence (2.3% wet wt day⁻¹); short day length seemed to be the major factor reducing growth rate during winter.
3. Mortality was close to zero during winter and 1.4—1.7% day⁻¹during other seasons. Total mortality from egg to adult was 99.6%.
4. The annual production was about 9g wet wt m⁻² year⁻¹ and the annual P/ ratio was 7.5. There was good agreement between the production values estimated by four methods. Production rate was highest in May (13 mg wet wt m⁻² day⁻¹) and zero in February.     

Global diversity of mayflies (Ephemeroptera, Insecta) in freshwater

The extant global Ephemeroptera fauna is represented by over 3,000 described species in 42 families and more than 400 genera. The highest generic diversity occurs in the Neotropics, with a correspondingly high species diversity, while the Palaearctic has the lowest generic diversity, but a high species diversity. Such distribution patterns may relate to how long evolutionary processes have been carrying on in isolation in a bioregion. Over an extended period, there may be extinction of species, but evolution of more genera. Dramatic extinction events such as the K-T mass extinction have affected current mayfly diversity and distribution. Climatic history plays an important role in the rate of speciation in an area, with regions which have been climatically stable over long periods having fewer species per genus, when compared to regions subjected to climatic stresses, such as glaciation. A total of 13 families are endemic to specific bioregions, with eight among them being monospecific. Most of these have restricted distributions which may be the result of them being the relict of a previously more diverse, but presently almost completely extinct family, or may be the consequence of vicariance events, resulting from evolution due to long-term isolation. Guest editors: E. V. Balian, C. Lévêque, H. Segers & K. Martens Freshwater Animal Diversity Assessment     

Patterns in the distribution of heptageniid (Ephemeroptera) species in Saskatchewan, Canada

SUMMARY. 1, The distribution patterns of twenty-one species of heptageniid mayflies were examined in Saskatchewan and west-central Manitoba, Canada. Presence/absence data from 174 stream sites were analysed using two complementary multivariate techniques, classification (Ward's method with Euclidean distance) and ordination (detrended correspondence analysis).
2. Multivariate analyses revealed two large, distinct groups of sites within the study area. The first included sites characterized by the presence of Heptagenia elegantula (Eaton) and Stenonema terminatum (Walsh). The second included sites characterized by the presence of Stenonema vicarium (Walker), Leucrocuta hebe (McDunnough) and/or Nixe species. The two groups contained, respectively, three and four subgroups.
3. Groups and subgroups defined by multivariate analyses were compared using large-scale environmental features obtained from topographic and other maps. Differences in environmental features between the two groups were mainly in stream order, terrestrial vegetation and annual number of degree-days (calculated from air temperature). The first group could be characterized as warm, large river habitat in grassland and parkland vegetation regions. The second included cooler, small to midsized streams and rivers in boreal forest. Most differences among subgroups were also in stream order, terrestrial vegetation and degree-days, although some subgroups in the second group differed also in stream gradient.
4. The observed groups and subgroups were not associated with drainage basins.     

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.     

Swarm behaviour and mate competition in mayflies (Ephemeroptera)

Although mayfly swarms are frequently cited as an example of lekking by insects, little is known about the behaviour of individuals within a swarm, or how mate-selection takes place. A study of five species of mayfly over a period of 10 consecutive years has revealed species-specific differences in the flight pattern of swarming males and in the ability of males to recognize swarms of their own species. Males of four of the five species jostle other males in the swarm at all times except when mating: mating pairs are not jostled. The pattern of jostling varies with the species. Measurements of the sperm content of the vesicula seminalis and of the wing length of members of individual swarms show that larger wing size is positively correlated with the presence of less sperm. The vesicula seminalis is always filled with sperm at the beginning of the imaginal stage and the testes regress before the beginning of the imaginal stage. If the volume of sperm in the vesicula seminalis is a valid index of mating success then males with larger wings have the highest success. Large wings may bestow an advantage during jostling. The males of Ephemera danica , which do not jostle, glide with outspread wings; these outspread wings may attract females, the largest wings being the most attractive. Females of all five species enter the swarm a few at a time, although many females may be resting beneath the swarm. This phased entry may decrease the attraction of the swarm for predators. The number of females in a swarm is not correlated with swarm size, and the factors which enable females to regulate their entry into a swarm remain obscure.     

Phylogeny of Ephemeroptera (mayflies) based on molecular evidence

This study represents the first molecular phylogeny for the Order Ephemeroptera. The analyses included 31 of the 37 families, representing approximately 24% of the genera. Fifteen families were supported as being monophyletic, five families were supported as nonmonophyletic, and 11 families were only represented by one species, and monophyly was not testable. The suborders Furcatergalia and Carapacea were supported as monophyletic while Setisura and Pisciforma were not supported as monophyletic. The superfamilies Ephemerelloidea and Caenoidea were supported as monophyletic while Baetoidea, Siphlonuroidea, Ephemeroidea, and Heptagenioidea were not. Baetidae was recovered as sister to the remaining clades. The mayfly gill to wing origin hypothesis was not supported nor refuted by these data. Mandibular tusks were supported as having at least one loss in Behningiidae and, together with the burrowing lifestyle, possibly two origins. The fishlike body form was supported as plesiomorphic for mayflies with multiple secondary losses. Topological sensitivity analysis was used as a tool to examine patterns concerning the stability of relationships across a parameter landscape, providing additional information that may not have been acquired otherwise.     

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.     

Integrative taxonomic revision of Campylocia (mayflies: Ephemeroptera,Euthyplociidae)

A morphological comparison of type and non-type material of species of Campylocia, including their junior synonyms, was conducted, in addition to neighbour joining based on K2P distances and Bayesian inference analyses of 376?bp of the mitochondrial gene cytochrome oxidase I (COI) of recently collected specimens. Results revealed the lack of distinguishing characters between C. bocainensis and C. dochmia supported by the molecular analysis, where the overlap of intra- and interspecific genetic divergences suggested genetic flow among individuals. Campylocia burmeisteri is revalidated as a senior synonym of Brazilian south-eastern species C. bocainensis and C. dochmia and of E. guntheri, formerly a synonym of C. anceps. Campylocia burmeisteri is redescribed based on material from its type-locality, Nova Friburgo, Rio de Janeiro State. Two new species, C. demoulini sp. nov. and C. araca sp. nov., are described from the Amazon rain forest and a third species, C. orosi sp. nov., is described from Costa Rica. Possible cryptic species and the utility of egg morphology in the taxonomy of Campylocia are discussed for the first time for the genus. A key to the identification of adult stages of Campylocia is provided based on male genitalia and egg morphologyhttp://zoobank.org/urn:lsid:zoobank.org:pub:6779DC2C-DB98-41DF-8C52-FF97366AAF7A     

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.     

Croatian mayflies (Insecta,Ephemeroptera): species diversity and distribution patterns

Knowledge of the mayfly biodiversity in the Balkan Peninsula is still far from complete. Compared to the neighbouring countries, the mayfly fauna in Croatia is very poorly known. Situated at the crossroads of central and Mediterranean Europe and the Balkan Peninsula, Croatia is divided into two ecoregions: Dinaric western Balkan and Pannonian lowland. Mayflies were sampled between 2003 and 2013 at 171 sites, and a total of 66 species was recorded. Combined with the literature data, the Croatian mayfly fauna reached a total of 79 taxa. Of these, 29 species were recorded for the first time in Croatia while 15 species were not previously recorded in Dinaric western Balkan ecoregion. Based on the mayfly assemblage, sampling sites were first structured by ecoregion and then by habitat type. In comparison with the surrounding countries, the Croatian mayfly fauna is the most similar to the Hungarian and Bosnian fauna. Some morphologically interesting taxa such as Baetis cf. nubecularis Eaton, 1898 and Rhithrogena from the diaphana group were recorded. Ephemera cf. parnassiana Demoulin, 1958, the species previously recorded only from Greece, was also recorded.     

Sense organs on the antennal flagellum of mayflies (Ephemeroptera)

A few very small sensory pegs with the characteristics of chemoreceptors are present in both males and females on the antennal flagellum of the mayflies, Stenacron interpunctatum (Say)

1 Stenacron interpunctatum (Say) is Stenonema interpunctatum (Say) of earlier literature.

and Cloeon sp. They are situated near the proximal end of the flagellum on its ventro-lateral surface and have escaped the attention of earlier investigators.