Size-related shifts in the physical habitat of two mayfly species (Ephemeroptera)

1. The spatial niche shifts of the nymphs of two mayfly species (Rhithrogena semicolorata and Ecdyonurus sp, gr. venosus) in relation to their growth were investigated in two lowland springs of northern Italy where they are dominant species. 2. Ordination of the different size classes of the two species revealed a canonical trend related to season and bottom roughness. 3. Larger nymphs of both species colonized rougher substrata. Differences between species in the roughness of areas colonized by small nymphs, which were very numerous, may have been due to competition for space.     

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.     

Automictic parthenogenesis of a geographically parthenogenetic mayfly,Ephoron shigae (Insecta: Ephemeroptera,Polymitarcyidae)

In the geographically parthenogenetic mayfly, Ephoron shigae, egg maturation and counts of chromosome number of unfertilized, parthenogenetic eggs were studied, in comparison with fertilized eggs from a bisexual population. The primary oocyte becomes mature through two successive maturation divisions. The first maturation division (meiotic division) takes place in the primary oocyte to produce a secondary oocyte and a first polar body. The second maturation division soon occurs in the secondary oocyte, in which the nucleus is divided into a mature egg nucleus (female pronucleus) and second polar body nuclei. The first polar body, in some cases, was successively divided into two polar bodies; in other instances, it was not divided. After the successive maturation division, the egg nucleus and the sister second polar body nucleus drew near to fuse into the zygote nucleus. The chromosome number was doubled in the zygote, and this conjugation initiates subsequent embryonic development. This suggests that, in E. shigae, the process of parthenogenetic recovery of diploidy is the automictic type categorized as the ‘terminal fusion’ pattern. © 2009 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 99 , 335–343.     

The daily activity patterns of mayfly nymphs (Ephemeroptera)

Nymphs of Baëtis rhodani (Pictet), Ephemerella ignita (Poda), Ecdyonurus venosus (Fabricius), Rhithrogena semicolorata (Curtis), and Heptagenia lateralis (Curtis) showed a similar endogenous rhythm of activity under conditions of natural illumination, continuous light and continuous darkness; and were most active at night when they moved from the lower to the upper surface of the stone. The nymphs were positively thigmotactic and negatively phototactic in flowing water, but these taxes and the endogenous activity rhythm of all species except Heptagenia lateralis changed markedly when the flow of water ceased.     

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.     

The larvae of the burrowing mayfly genus Tortopus (Ephemeroptera: Polymitarcyidae)

The larval stage of Tortopus is redescribed based on three species: T. puella from North America, the only species of the genus previously known from larva, and the larvae of T. obscuripennis and T. sarae from South America described here. Generic characters of the larva include: relatively large finger-like gill near base of maxilla, inner margin of mandibular tusks with a subdistal tubercle, straight or weakly convex frontal ridge present between antennae, reduced unilamellated gill on abdominal segment I. Additionally the male imagines of both Neotropical species are described for the first time, and T. obscuripennis is recorded from Bolivia. Diagnoses, SEM photographs, and illustrations are given for the new stages described and for the identification of the three Tortopus species known as larvae.     

Potential for parthenogenesis of virgin females in a bisexual population of the geographically parthenogenetic mayfly Ephoron shigae (Insecta: Ephemeroptera,Polymitarcyidae)

The burrowing polymitarcyid mayfly Ephoron shigae is a geographically parthenogenetic species. Interestingly, the distributions of the bisexual and unisexual populations overlap broadly in their respective geographic ranges. In this mayfly, obligatory diploid thelytoky appears within unisexual populations. In the present study, we examined the potential for parthenogenesis or the parthenogenetic ability of females in a bisexual population aiming to understand the emergence of unisexual populations. The results obtained revealed that females in the examined bisexual populations showed a potential for diploid thelytoky as also seen in the unisexual populations, although, in females from bisexual populations, the development success rates of their unfertilized eggs were considerably lower than those of virgin females from unisexual populations. In the three bisexual reproducing species (Ephemera japonica, Ephemera strigata, and Ephemera orientalis) in the closely‐related family Ephemeridae, diploid thelytoky (i.e. tychoparthenogenesis; < 3%) was also observed. However, in this case, the parthenogenetic development success rates of unfertilized eggs were significantly lower than those of virgin females in the bisexual (Hino‐yosui Irrigation Canal) population of E. shigae. Accordingly, we suggest that parthenogenetic ability (i.e. tychoparthenogenesis or facultative parthenogenesis) in bisexual populations of E. shigae may facilitate the evolutionary transition to unisexual populations with fully obligatory parthenogenesis. © 2009 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 99 , 326–334.     

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.     

Phylogeny of the mayfly family Leptohyphidae (Insecta: Ephemeroptera) in South America

Abstract.  A cladistic analysis of the South American members of the Ephemeropteran family Leptohyphidae is presented. A matrix of 73 taxa and 124 morphological characters was analysed under two distinct weighting criteria (implied weighting, which weights characters as a whole, and self-weighted optimization, which differentially weights character state transformations). To assess the monophyly of the Leptohyphidae, representatives of Ephemerellidae, Ephemerythidae, Machadorythidae, Teloganodidae, Tricorythidae, Coryphoridae and Melanemerellidae were also included. Trees were rooted in Ephemerellidae. Conspicuous differences in consensus topology occur when transformation costs among character states are weighted (including asymmetries). The differences in the assessments of character reliability in the two weighting criteria used are discussed. In many cases, self-weighting, in allowing for asymmetries in transformation costs, considered many of the character state transformations as more reliable (= informative) than implied weights (which needlessly down-weighted the whole character). The results confirm the monophyly of Leptohyphidae and support its sister-group relationship with Coryphoridae. The shortest trees do not support the recently proposed division of Leptohyphidae into two subfamilies. Ephemerelloidea higher classification is discussed briefly.     

Comparison of mayfly (Ephemeroptera) taxocenes of permanent and intermittent Central European small streams via species traits

The recurrent drying out of small streams in past decades has shown an urgent need to pay attention to the impact of global climate change. The objectives of this study were to describe the effect of drying out on the composition of the mayfly taxocene and evaluate the relevance of individual species traits for survival of mayflies to drying out. The mayfly taxocenes of two model localities, one at an intermittent and one at a permanent brook, were investigated in 2002, 2003 and 2005. Compared with the permanent stream, the taxocene of the intermittent stream was short of nine species, foremost rheobionts and high oxygen demand species. To explain further differences between both stream types in survival and recolonisation ability, 15 species traits were evaluated. These included so-called “ecological traits” (e.g., habitat and substrate range, density, distribution, current velocity adaptation) and “biological traits” connected with life cycle and larval/adult adaptations. Species showing the highest number of advantageous traits (with only exception of Electrogena sp. cf. ujhelyii — species of taxonomically unclear status) were able to successfully survive under the unfavourable conditions of the intermittent brook. Biological traits considered more important in many respects seem to be good predictors for assessing sensitivity to extreme temperature changes, hydrological regime fluctuations and the survival/recolonisation ability of species in exposed habitats.     

Biometric studies of some stoneflies and a mayfly (Plecoptera and Ephemeroptera)

Aspects of the nymphal/adult developmental change were investigated in biometric studies of several species of Plecoptera: Nemouridae near Schlitz, Hesse, Germany. Preliminary information on the mayfly, Baetis vernus Curtis, is also provided. Nemourid nymphs pass through 3 wing bearing stages before reaching adulthood. Instars can be identified by their characteristic shapes, as expressed by the wing length/head width (WL/HW) ratio. Size does not allow instar discrimination, mainly due to sexual size differences. HW is ca 10% larger in last instar female than in male nemourid nymphs; exuviae shed at the moult to adult represent about 14% of nymphal ash free dry weight (AFDW). Biomass lost with exuviae during the many larval moults should be accounted for in estimates of production. Freshly emerged nemourid females are about 6% larger and 30% heavier than males. The HW/AFDW relationship is the same in both sexes. Through terrestrial feeding during adult life, males double their weight on average. Mature females are up to three times heavier than freshly emerged ones. They invest about 30% of their final AFDW in reproduction.Shape of last instar nymphal Baetis was expressed as the ratio wing length/mesonotum length. It is size-dependent, a characteristic, instar-specific shape may not occur in this mayfly. Nymphal and subimaginal exuviae together represent about 14% of last instar nymphal dry weight. Females of Baetis are about 55% heavier than males. Unlike in Plecoptera, the size/weight (ML/AFDW) relationship differs between sexes.     

Molecular data and species distribution models reveal the Pleistocene history of the mayfly Ameletus inopinatus (Ephemeroptera: Siphlonuridae)1

1. We investigated the Pleistocene and Holocene history of the rare mayfly Ameletus inopinatus EATON 1887 (Ephemeroptera: Siphlonuridae) in Europe. We used A. inopinatus as a model species to explore the phylogeography of montane, cold‐tolerant aquatic insects with arctic–alpine distributions. 2. Using species distribution models, we developed hypotheses about the species demographic history in Central Europe and the recolonisation history of Fennoscandia. We tested these hypotheses using mitochondrial cytochrome oxidase I (mtCOI) sequence data and compared our genetic results with previously generated microsatellite data to explore genetic diversity distributions of A. inopinatus. 3. We observed old lineages, deep splits and almost complete lineage sorting of mtCOI sequences among mountain ranges. These results support a periglacial survival, i.e. persistence at the periphery of Pleistocene glaciers in Central Europe. 4. There was strong differentiation between the Fennoscandian and all other populations, indicating that Fennoscandia was recolonised from a refugium not accounted for in our sampling. High degrees of population genetic structure within the northern samples suggest that Fennoscandia was recolonised by more than one lineage. However, this structure was not apparent in previously published microsatellite data, consistent with secondary contact without sexual incompatibility or with sex‐biased dispersal. 5. Our demographic analyses indicate that (i) the separation of northern and Central European lineages occurred during the early Pleistocene; (ii) Central European populations have persisted independently throughout the Pleistocene and (iii) the species extended its range about 150 000 years ago.     

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.

     

Ultrastructure of the antennal sensilla of the mayfly Baetis rhodani (Pictet) (Ephemeroptera : Baetidae)

The distribution and fine morphology of antennal sensilla of nymphal and adult mayfly, Baetis rhodani (Ephemeroptera : Baetidae), were examined. In the nymph, various kinds of sensilla (chaetica, basiconica, coeloconica and cuticular pits) are differently arranged on the antennal segments, whereas sensilla campaniformia delimit the distal border of the pedicel. A peculiar kind of sensillum basiconicum, named flat-tipped sensillum, is present along the entire antenna, even though in the flagellum it has a regular arrangement between the cuticular lobes that delimit the distal border of each article. In the subimago the scape and pedicel are profusely covered with microtrichia and scattered sensilla trichodea, whereas the flagellum shows cuticular ribs. Sensilla coeloconica are present along the ventral side of the flagellum. In the imago, the antenna is completely decorated with scales among which sensilla trichodea and sensilla coeloconica occasionally occur. As in the nymph, adult mayflies have a ring of sensilla campaniformia along the distal border of the pedicel. When compared with nymphal antennae, those of adults have fewer types of sensilla, presumably in relation to the short, non-feeding terrestrial life.     

New mayfly nymphs (Insecta: Ephemeroptera) from Yixian Formation,China

Three new species of mayfly nymphs, Clavineta excavata sp. nov., Clavineta brevinodia sp. nov., and Siberiogenites branchicillus sp. nov., from the Yixian Formation (Late Jurassic-Early Cretaceous) in western Liaoning Province, China, are described. These mature nymphs are small-sized, with the body less than 16.0 mm long. The systematic position of the genera Clavineta and Siberiogenites, which are recorded for the first time in the Yixian Formation, is briefly discussed.     

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.     

Habitat characterization of the morphologically similar mayfly larvae,Caenis and Tricorythodes (Ephemeroptera)

The larvae of Caenis (Caenidae) and Tricorythodes (Tricorythidae), once considered to be confamilial, have notable morphological and behavioural similarities. Univariate and multivariate (discriminant analysis) techniques were used to determine which environmental variables best characterized the larval habitats of Caenis and Tricorythodes at 40 sample sites on 29 rivers within the Interior Plains of Alberta, Canada. River width, depth and substrate type distinguished riverine habitats of the two genera. Larvae of Tricorythodes occurred in wide rivers of varying depths that possessed coarse substrates. Although larvae of Caenis occurred in a variety of habitats, they were found more frequently on stable substrates in narrow, deep rivers.