Influence of morphohydraulic habitat structure on invertebrate communities (Ephemeroptera,Plecoptera and Trichoptera)

Fluvial geomorphology proposes the methodology of cognition and assessment of the riverine landscape and points to the possibilities of exploitation of its results in hydrobiological research. Habitat structure of two reaches of the Drietomica brook (Biele Karpaty Mts, Slovakia) was assesed at level of morphological and morphohydraulic units in the sense of the River Morphology Hierarchical Classification (RMHC)). Physical habitats were described by flow hydraulics and substrate properties as directly measured variables (current velocity, depth, substrate size) and related variables (flow type, Froude and Reynolds numbers). According to the shear stress (expressed by Fr and Re), the morphological units were divided into two main groups — with low shear stress — pools, glides, edgewaters, bar nooks and bars; with high shear stress — riffles, runs, rapids and scours; characterized also by different Ephemeroptera, Plecoptera and Trichoptera (EPT) communities. The EPT communities were analyzed in relation to the morphological, hydraulic and substrate characteristics of the stream channel. The main environmental gradient responsible for the variation in EPT fauna was found using Principal Component Analysis and was related to gradient of flow in term of current velocity and other hydraulic attributes covered by Fr and Re numbers. The EPT communities (by means of abundance, feeding types, current, microhabitat and zonation preferences) showed preferences for different morphological units, flow type and current velocity. Depth and substrate grain size showed only weak relation to EPT communities.     

A DNA barcode library for Germany′s mayflies,stoneflies and caddisflies (Ephemeroptera,Plecoptera and Trichoptera)

Mayflies, stoneflies and caddisflies (Ephemeroptera, Plecoptera and Trichoptera) are prominent representatives of aquatic macroinvertebrates, commonly used as indicator organisms for water quality and ecosystem assessments. However, unambiguous morphological identification of EPT species, especially their immature life stages, is a challenging, yet fundamental task. A comprehensive DNA barcode library based upon taxonomically well‐curated specimens is needed to overcome the problematic identification. Once available, this library will support the implementation of fast, cost‐efficient and reliable DNA‐based identifications and assessments of ecological status. This study represents a major step towards a DNA barcode reference library as it covers for two‐thirds of Germany's EPT species including 2,613 individuals belonging to 363 identified species. As such, it provides coverage for 38 of 44 families (86%) and practically all major bioindicator species. DNA barcode compliant sequences (≥500 bp) were recovered from 98.74% of the analysed specimens. Whereas most species (325, i.e., 89.53%) were unambiguously assigned to a single Barcode Index Number (BIN) by its COI sequence, 38 species (18 Ephemeroptera, nine Plecoptera and 11 Trichoptera) were assigned to a total of 89 BINs. Most of these additional BINs formed nearest neighbour clusters, reflecting the discrimination of geographical subclades of a currently recognized species. BIN sharing was uncommon, involving only two species pairs of Ephemeroptera. Interestingly, both maximum pairwise and nearest neighbour distances were substantially higher for Ephemeroptera compared to Plecoptera and Trichoptera, possibly indicating older speciation events, stronger positive selection or faster rate of molecular evolution.     

DNA barcoding of stoneflies (Plecoptera) in a general genetics course

DNA Barcoding is a powerful molecular biology tool for the identification of species, analysis of gene flow from within and between populations and evaluating species concepts. The process can be incorporated into a college genetics or bioinformatics core curriculum. Here we demonstrate how, by simply using Plecoptera, we actively engaged students in DNA extraction, amplification and DNA Barcoding. As a result, 38 new DNA sequences were accepted by an international DNA database, with potentially three of them being new species to the database. More importantly, through multiple assessment measurements, students showed a high degree of learning took place and that all student learning objectives were met.     

Global diversity of stoneflies (Plecoptera; Insecta) in freshwater

Plecoptera, or stoneflies, is a small order of hemimetabolous insects: according to our data, more than 3,497 species have been described so far in the world. The total number of species has enormously increased in the last 30 years (2,000 species estimated in 1976) and, if the trend continues, then it will nearly double in the near future. The order is divided into the suborders Arctoperlaria and Antarctoperlaria, and includes 16 families: 12 arctoperlarian and 4 antarctoperlarian. The Arctoperlaria account for a total number of 3,179 species, and Antarctoperlaria, only 318 species. The total number of genera is 286. We give in this article the estimated number of species for each family. The fauna and diversity of stonefly in North America (650 species reported) and Europe (426 species) are best known. Nevertheless, in the last 25 years, a mean of 2.6 Plecoptera species per year were described in Europe. Stonefly-faunas of Australia (191 species, Tasmania included) and New Zealand (104 species) are relatively well-known, while our knowledge of the Plecoptera of Central and South America (95 and 378 species respectively) is poor and still not representative of the real diversity. Africa has a reduced stonefly fauna (126 species). Asian stonefly diversity (approximately 1,527 species) is much greater than that of Europe or North America despite the fact that, except for Japan and Asiatic Russia that have been well studied, our knowledge of the remaining Asiatic areas is extremely poor. Even though our data indicate the Holarctic Region as the diversity hot-spot for the order, the analysis of the specific diversity divided by family suggests also an important role of tropical stoneflies. Guest editors: E.V. Balian, C. Lévêque, H. Segers & K. Martens Freshwater Animal Diversity Assessment     

Post-emergence migration of stoneflies (Plecoptera) into the nearby forest

Abstract. 1. The post-emergence lateral migration of both sexes of eight stonefly species was examined in a dystrophic, fourth-order forest river in eastern Finland.
2. For this purpose, 7351 stonefly adults were collected with eighteen trunk funnels positioned in rows of six at distances of approximately 1, 15 and 60 m from the river. A further 1880 adults were also caught from the vegetation of the bank zone by sweep netting and with slit traps.
3. The species could be grouped into two types with respect to migration distance: Isoperla difformis, I.grammatica and Leuctra fusca tended to stay in the bank area, whereas Nemoura flexuosa, N.avicularis, Amphinemura borealis, L.hippopus and N.cinerea tended to disperse into the forest, so that the majority were found some distance away from the shore.
4. The males of the leuctrids, N.flexuosa and N.cinerea migrated farther than the females.
5. The sex ratio was significantly biased in all species except N.flexuosa. The isoperlids, leuctrids and A.borealis showed a significant predominance of females in the trunk funnel catches, but males were significantly dominant in N.avicularis and N.cinerea. The material caught by other methods reversed the ratio for I.difformis and N.avicularis. Comparison of the sex ratios of the species with other reports revealed marked variation and deviation from unity, much of which could be attributed to bias introduced by the sampling methods.
6. Lateral migration seems to be the first phase in the colonization cycle, although the latter as such was not studied here. Adults of Euholognatha species migrate farther than those of Systellognatha, a difference of which may be due to their ability to feed as adults.     

Egg characteristics and oviposition behaviour of the aquatic insect orders Ephemeroptera,Plecoptera and Trichoptera in New Zealand: a review

The abundance and distribution of aquatic insects may be controlled by factors at any stage of the life cycle. To understand the potential role of factors affecting egg and adult stages and oviposition, basic information on egg characteristics and oviposition behaviour is required. However, published information on New Zealand Ephemeroptera, Plecoptera and Trichoptera (EPT) is scattered and very limited. This review compiles current knowledge of New Zealand EPT egg characteristics and oviposition behaviour using published information and unpublished observations. Where direct observations are lacking, information on closely related species overseas, and inferences based on female morphology, are included. Eggs show a variety of physical characteristics that may influence egg retention, transport and distribution. Oviposition behaviour is highly selective among some taxa, but appears generalist among others, though the latter may still require specific cues. Identifying and providing for adult, egg and oviposition requirements may be fundamental to re-establishing EPT populations in restored aquatic habitats.     

Microsatellite loci for the mayfly Baetis rhodani (Baetidae,Ephemeroptera)

The first microsatellite primers are described for the mayfly family Baetidae (order Ephemeroptera). Seven polymorphic loci were isolated and characterized for the species Baetis rhodani from two enriched genomic libraries. A total of 183 individuals were genotyped using these loci. Allelic diversity was high for all loci, and observed heterozygosities ranged between 0.382 and 0.772. A heterozygote deficiency was detected in some loci, suggesting the presence of ‘null’ alleles.     

Relationship between benthic insects (Ephemeroptera, Plecoptera, Coleoptera, Trichoptera) and temperature in small and medium-sized streams in Germany: A multivariate study

Benthic insect communities (Ephemeroptera, Plecoptera, Coleoptera, Trichoptera) were studied together with water temperature and environmental parameters in streams between June 2000 and June 2001. The sampling area consisted of 20 sites in small and medium-sized streams located in the lower mountainous area of Central Europe. Temperature was recorded nearly continuously and several physicochemical and environmental variables were assessed. Macroinvertebrates were sampled both in spring and summer. Data-sets of species abundance and occurrence were analysed using multivariate techniques and were correlated to the thermal and environmental conditions of the streams. The temperature preferences of the species were compared to published data-sets on their autecological characteristics. Up to 29% of the variability in the Ephemeroptera, Plecoptera, Trichoptera and Coleoptera community was explained by summer temperature variation in the data-sets for both small and medium-sized streams. A smaller, but significant part of the variability in species distribution was explained by conductivity, substratum type, and the percent coverage of local riparian forest. Compared to small streams, temperature was less important for the macroinvertebrate composition in medium-sized streams. This result is likely due to the more tolerant, eurythermic species composition in larger streams. A total of 33 Ephemeroptera, Plecoptera, Coleoptera and Trichoptera taxa were positively correlated and 28 taxa were negatively correlated to summer temperature patterns. The temperature preferences of taxa considered in this study were related to species traits, such as egg dormancies and life cycle plasticity.     

Dispersal,feeding and parasitism of adult stoneflies (Plecoptera) at a New Zealand forest stream

The temporal occurrence and dispersal of stoneflies were investigated at a forest stream in the South Island, New Zealand. Although 12 species were taken in Malaise traps, only Spaniocerca zelandica and Cristaperla fimbria (Notonemouridae) were abundant. Adults of S. zelandica were present throughout the year and a wide range of different-sized nymphs, including very small individuals, was also found in all months. In contrast, the emergence period of C. fimbria was limited to about four months in summer and as nymphs are mainly hyporheic, few were taken in Surber samples. Malaise trapping on a 16 m long transect away from the stream and with more distantly located traps in forest and grassland resulted in a very high proportion of males and females of both species being caught within 1 m of the stream edge. Captured females of both species included immature, gravid, and spent individuals. Males and females of both species had almost identical diets dominated by sooty mould fungi, fungal spores, pollen and fine particulate organic matter. High proportions of them were also infected by the encysted larvae of a gordian worm (Nematomorpha) and may be important vectors in its life cycle.     

Phenology and Abundance of Ephemeroptera,Plecoptera and Trichoptera Caught by Emergence Traps at the Weidlingbach near Vienna,Austria

A total of 1136 Ephemeroptera, Plecoptera and Trichoptera were caught from March 1993 to March 1994 at the Weidlingbach, a first to fourth order forest brook near Vienna. Austria, using pyramid-type emergence traps at tree sampling stations (390, 360 and 260 m above sea level). Five species comprised 67.1 % of the catch: the ephemeropterans Paraleptophlebia submarginata (Stephens), Habroleptoides confusa Sartori and Jacob and Habrophlebia lauta Eaton, and the plecopterans Nemoura flexuosa Aubert and Leuctra hippopus Kempny. Most taxa emerged from April to August, but the plecopterans Leuctra prima Kempny and Capnia bifrons (Newman) were caught early in the year (January to April), and Leuctra digitata Kempny (Plecoptera), Chaetopteryx fusca Brauer and Chaetopteryx major McLachlan (Trichoptera) emerged in September and October. In Ephemeroptera and Trichoptera, similarities between species groups were highest at the two upstream sampling stations; in Plecoptera and in pooled taxa, however, similarities were highest at the middle and downstream sites. Except in Trichoptera, diversity was highest at the middle station situated at 360 m above sea level.     

New mayfly nymphs from the Jurassic of Northern and Northeastern China (Insecta: Ephemeroptera)

Four new species of mayfly nymphs, Furvoneta relicta sp. nov., Clavineta eximia sp. nov. (originally Mesoneta antiqua), Mesobaetis latifilamentacea sp. nov., and Stackelbergisca cylindrata sp. nov. are described from the Jurassic of China. All these species may have inhabited lakes rather than flowing water. The geological age and stratigraphical correlation of the nymph-bearing strata are briefly reviewed.     

The effects of extended photoperiods on the drift of the mayfly ephemerella subvaria McDunnough (Ephemeroptera : Ephemerellidae)

Field drift studies indicated that the nocturnal drift density of E. subvaria nymphs was greater in early May than in early November.Laboratory studies showed that the number of individuals appearing in the drift was a linear function of the duration of the preceding photoperiod. Nymphs had a greater propensity to drift when they were not in a state of active growth than when they were growing. The tendency of individuals in a single laboratory population to drift was observed to change under conditions of constant temperature and randomized photoperiod. This suggests that the shift was due to some internal physiological change rather than to an external cue.It is suggested that drift in E. subvaria functions as a method relocation from fast-water areas to slow-water pools and stream margins. Redistribution to these areas may reduce mortality incurred during spring run-off and during emergence.     

Structure and spatial variability of mayfly (Ephemeroptera) communities in the upper Hron River basin

Mayflies were sampled from eleven sites in six tributaries of the upper Hron River. Mayfly communities were ordinated and related to environmental factors using multiple regression and principal component analysis. Communities of headwater sites were dominated by Rhithrogena iridina which was gradually replaced by Baetis alpinus, at the lower reaches. Total mayfly species richness was inversely related to stream slope and overall abundance was inversely related to altitude. The prominent factor influencing mayfly communities was a complex longitudinal gradient as proposed by the river continuum concept. There were no significant differences in community composition and structure between the left-and right-hand tributaries of the Hron River.     

Discovery of an alien species of mayfly in South America (Ephemeroptera)

Despite its wide, almost worldwide distribution, the mayfly genus Cloeon Leach, 1815 (Ephemeroptera: Baetidae) is restricted in the Western hemisphere to North America, where a single species is reported. In the Neotropics, except for some species wrongly attributed to the genus in the past, there are no records of Cloeon. Recently, however, specimens of true Cloeon were collected along the coast of Espírito Santo, Southeastern Brazil. In order to verify the hypothesis that this species was recently introduced to Brazil, our aim was to identify the species based on morphological and molecular characters and to confirm the presence of true representatives of the genus in the Neotropics. Our results revealed that the specimens found in Brazil belong to the Afrotropical species C. smaeleni Lestage, 1924. The identity of the species, its distribution, along with its previous absence in regularly sampled sites, is a clear sign that the specimens of C. smaeleni found in Espírito Santo are introduced, well established, and that the colonization took place very recently.     

Species‐specific communication bars interspecific mating between syntopic species of Zwicknia stoneflies (Plecoptera: Capniidae)

Most Northern Hemisphere stoneflies have species‐specific mating signals that are generally thought to constitute a barrier against interspecific mating. We tested this hypothesis in two species of the genus Zwicknia that have only very recently been recognised as distinct species, and that were found to occur together in a stream in Lower Saxony, Germany. Analyses of molecular markers COI and 28S in combination with wing length (distinguishing males of both species) and mating signals revealed no instance of hybridisation among 23 studied specimens. In addition, eleven further males identified on the basis of morphology alone all produced the expected species‐specific signal. Females and males of both species were presented with played back conspecific and heterospecific signals and duetting sequences, and responded only to conspecific stimuli. This lends support to the hypothesis that the intersexual communication system functions as an important pre‐mating barrier against gene flow, although post‐mating isolation cannot be excluded. Interspecific mating did occur when a mixed pair was confined together in a small container. Males of both species were found to call in response to played back duetting sequences with stereotypic latencies that are clearly longer than the latencies in male‐female duets. We interpret this as an indication of eavesdropping behaviour coupled with attempts to take over the perceived duet. © 2014 The Linnean Society of London, Biological Journal of the Linnean Society, 2014, 113 , 969–980.     

Resource division by perlid stoneflies (Plecoptera) in a lake outlet ecosystem

In a stream draining a productive lake, two abundant (250/m²) stoneflies Calineuria californica and Hesperoperla pacifica differed in population size structure, habitat use and food although overlap in resource use was extensive. Life cycles of two (Calineuria) and three years (Hesperoperla) were indicated. A multivariate analysis of habitat use demonstrated size dependent habitat selection in both species and between species habitat differences. Feeding habits differed although only the largest Hesperoperla had a food refuge. Diets of individual nymphs represented an interaction of species, size and habitat.     

Inter- and intra-specific differences in the number of larval instars in British populations of 24 species of stoneflies (Plecoptera)

1. Ontogenetic changes during the life cycle of aquatic insects are important not only in life‐history studies but also in evaluating food‐web structure. They require information on the growth and number of larval instars but such information is lacking for many species, including Plecoptera. Therefore, the chief objectives of the present study were to determine inter‐ and intra‐specific differences in the number of larval instars in British populations of 24 species of stoneflies, to test Dyar’s hypothesis that growth followed a geometric progression, and to synthesise this information with previously published values for four British species. 2. Larvae were reared at constant temperatures in the laboratory from eggs from 63 populations (one to six populations per species). First instars from each population were divided into three batches and each batch was reared at one of three constant temperatures. For each species, the rearing temperature and source population had no significant effect on the mean size of each larval instar. 3. The relationship between the geometric mean length of each instar and instar number was well described by an exponential equation (P < 0.001, r² > 0.9 for all species), thus supporting Dyar’s hypothesis. Only one species, Brachyptera risi, had the same number of instars for males and females (12–13). For the other 15 herbivorous species and the four smaller carnivorous ones, the number of instars was higher for females than males (range 11–16 for males, 12–17 for females). The larger size of the females was due to their additional instars, not a sex difference in growth rates. In contrast, there was a clear growth separation of the sexes after the 9th or 10th instar for the four largest carnivores. The number of larval instars was highest for these four species (range 16–19 for males, 18–23 for females), and females were much larger than males. 4. A multiple regression equation with data from the present and previous studies (n = 27) showed that variability in the mean length of the first instar and the maximum number of larval instars for each species accounted for 88% and 91% of the variability in the mean length of the final instar for males and females, respectively. 5. Values for Plecoptera in other countries were in general agreement with those in the present study, especially in the same families. Two old, but widely quoted, high values are doubtful. The present study and four previous ones provide a sound basis for ontogenetic studies on 28 species of Plecoptera and their role in aquatic ecosystems.     

The oxygen consumption of mayfly (Ephemeroptera) and stonefly (Plecoptera) larvae at different oxygen concentration

1. The aim of this investigation was to elucidate how four acquatic insect larvae, from different habitats and having different respiratory organs or types of respiratory regulation, react to a lowered oxygen concentration, and how their oxygen consumption is affected. The species investigated were the stoneflies Taeniopteryx nebulosa, Diura nanseni and Nemoura cincerea and the mayfly Cloëon dipterum.
2. The measurements were performed in a respiratory apparatus of open, flowing-water type. Its design is shown in Fig. 1. Water of known oxygen concentration was allowed to flow past the experimental larvae. The oxygen consumption of the larvae was calculated from the lowering of the oxygen concentration in which ensued.
3. The water used in the experiments was standardized, so that the electrode had the necessary stability (conductivity 470 micromhos/cm). The calcium ion was excluded in order to prevent the precipitation of CaCO₃ in the electrode capillary.
4. A large variation in the values of oxygen consumption was found as seen in Fig. 2–5. The reason for that is a corresponding variation in the motor activity of the experimental animals.
5. The physiological reasons for the general form of the curves A and C in Fig. 2–5 are discussed. The curves A and C represent oxygen consumption of the larvae at different degrees of stimulation, entailing different levels of motor activity. Curve A represents intentinally activated animals, curve C non-activated, motionless animals. The curves A and C are boundary curves corresponding to a sort of scope for activity of the animals. Over this scope area a series of curves of the same form could in principal be construed, representing different degrees of stimulation.
6. Within a certain oxygen concentration interval a motor activation was observed caused by a reduced oxygen concentration. The result of that activation can be seen in Fig. 2–5 as a zone with no or very few oxygen consumption values between curve C and D. The more easily activated the species is, the broader the zone will be. Cloën has the most narrow zone and was observed to be less activated than the other species.
7. Small larvae of Cloën (2–4 mm and 42–6 mm) and Nemoura (2–4 mm) showed clearly a greater ability to take up oxygen at low oxygen concentrations than full-grown larvae (see Fig. 8 and 9).
8. The critical point on the curve representing mean oxygen consumption as a function of oxygen concentrations was found to be at 2–5 mg O₂/1 for Taeniopteryx and Diura, at 2.2–2.5 mg O₂/1 for Cloëon, and at about 2–7 mg O₂/1 for Nemoura. The values refer to 8°. Cloëon is the species which is exposed to the greatest variations in oxygen concentration in its natural environment.
9. No influence on the oxygen consumption of starvation for 4 to 5 days was found. No difference between the oxygen consumption values obtained in the presence or in the absence of calcium ions could be observed during the experiments (Fig. 10, 11).
10. The basic picture obtained in this investigation is a set of oxygen consumption values scattered between a curve connecting highest values obtained and a curve of the standard metabolism together with a zone in which the larvae are activated by reduced oxygen concentrations. This picture is presumed to be general in aquatic animals with a well developed motor activity.