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.     

Small-scale patterns of genetic variation in the mayfly Bungona narilla (Ephemeroptera: Baetidae) in rainforest streams, south-east Queensland

SUMMARY 1. Genetic structure of the mayfly Bungona narilla was examined using allozymes and a section of the cytochrome oxidase I gene.
2. The study had two major aims. The first was to determine whether patterns of genetic variation in mitochondrial DNA were similar to those found previously for allozymes, i.e. that more variation was evident among pools within a single stream than among streams. The mitochondrial DNA results were similar to those reported previously for allozymes, supporting the idea that larvae within any particular pool were unrepresentative of the total population and may result from a few matings.
3. The second aim was to test the hypothesis that the variation among pools within a stream was greater after dry periods than after wet periods. This was because after wet periods, larvae would have greater opportunity for mixing because of movement among pools. This hypothesis was partly supported by the mitochondrial DNA data but not by the allozyme data, in which variation among pools was extremely low on both sampling occasions. The reasons for this difference are unclear.     

Allozyme diversity and population genetic structure of the caddisfly Orthopsyche fimbriata and the mayfly Acanthophlebia cruentata in New Zealand streams

1. Allozymes were used to measure genetic variation within and among regional populations of the caddisfly Orthopsyche fimbriata and the mayfly Acanthophlebia cruentata in North Island New Zealand streams.
2. High levels of genetic differentiation were recorded in populations of O. fimbriata within and among catchments separated by more than 100 km, but little or no differentiation in populations separated by around 10 km. The Auckland isthmus appears to be a major barrier to north–south gene flow, with nearly fixed allelic differences at one locus. Genotype frequencies conformed to Hardy–Weinberg equilibrium.
3.  Acanthophlebia cruentata had low levels of genetic variation; the results are unexpected given that O. fimbriata apparently has greater potential for dispersal. The limited genetic data for A. cruentata provided evidence for genetic differentiation among populations separated by around 100 km, or more, within catchments and some differentiation between catchment populations separated by only 25 km.     

Egg envelopes of Baetis rhodani and Cloeon dipterum (Ephemeroptera, Baetidae): a comparative analysis between an oviparous and an ovoviviparous species

A comparative ultrastructural investigation on the eggshell (vitelline and chorionic envelopes) has been carried out in the nymphs of two mayfly species encompassed into the Baetidae, namely Baetis rhodani and Cloeon dipterum. During oocyte differentiation in the meroistic telotrophic ovarioles, gametes are connected to discrete nurse cells by trophic cords. In B. rhodani, each ovariole contains several eggs arranged in sequence, whereas in C. dipterum each contains a single egg. Follicle cells are competent for vitelline and chorionic envelope synthesis. Baetis rhodani is an oviparous species and the chorion is fairly thick, formed by an alveolate endochorion and a fibrillar exochorion delimited by a honey‐comb roof. Cloeon dipterum stands out among Ephemeroptera for being ovoviviparous. In B. rhodani, ovulation starts in the older nymphs with dark wing‐pads, whereas in C. dipterum choriogenesis ends in the imaginal stage. Here the chorion is very thin and laid eggs hatch almost immediately, allowing the larvulae to move out. The maturation of a single egg per ovariole is synchronized with the achievement of the adult stage. The acquisition of ovoviviparity has led to remarkable changes in the ovariole organization along with a simplification of the eggshell structure.     

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.     

Evolution and island endemism of morphologically cryptic Baetis and Cloeon species (Ephemeroptera,Baetidae) on the Canary Islands and Madeira

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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.     

Cryptic diversity and multiple origins of the widespread mayfly species group Baetis rhodani (Ephemeroptera: Baetidae) on northwestern Mediterranean islands

How the often highly endemic biodiversity of islands originated has been debated for decades, and it remains a fervid research ground. Here, using mitochondrial and nuclear gene sequence analyses, we investigate the diversity, phylogenetic relationships, and evolutionary history of the mayfly Baetis gr. rhodani on the three largest northwestern Mediterranean islands (Sardinia, Corsica, Elba). We identify three distinct, largely co‐distributed, and deeply differentiated lineages, with divergences tentatively dated back to the Eocene–Oligocene transition. Bayesian population structure analyses reveal a lack of gene exchange between them, even at sites where they are syntopic, indicating that these lineages belong to three putative species. Their phylogenetic relationships with continental relatives, together with the dating estimates, support a role for three processes contributing to this diversity: (1) vicariance, primed by microplate disjunction and oceanic transgression; (2) dispersal from the continent; and (3) speciation within the island group. Thus, our results do not point toward a prevailing role for any of the previously invoked processes. Rather, they suggest that a variety of processes equally contributed to shape the diverse and endemic biota of this group of islands.     

MtDNA COI barcodes reveal cryptic diversity in the Baetis vernus group (Ephemeroptera, Baetidae)

Partial mitochondrial COI sequences (barcoding fragment) were explored for the understanding of the species boundaries of Baetis vernus group taxa (Ephemeroptera, Baetidae) in northern Europe. We sampled all species of this group occurring in Finland, but focused on taxa for which morphological and taxonomical confusion have been most apparent. The sequence matrix comprised 627 nucleotides for 96 specimens, and was analysed using parsimony. Results provided strong evidence that Baetis macani Kimmins and B. vernus Curtis comprise morphologically cryptic but molecularly distinct taxa, as intraspecific uncorrected divergences within haplogroups ranged between 0.3% and 1.4% and interspecific divergences were from 13.1% to 16.5%. These interesting findings prompt for further taxonomic studies of B. vernus taxa using more extensive specimen sampling from the known distributional areas in the Palaearctic/Holarctic region for better understanding of haplotype distributions. We stress the importance of integration of morphological and molecular data, and the necessity to employ additional nuclear DNA sequence data.     

Genetic variation and social structure: a case-study from Africa

Understanding the relations between social and biological factors in determining the evolution of human populations is one of the principal aims of bio-anthropological research. The recent introduction of unilinearly transmitted polymorphisms of mitochondrial DNA and Y-chromosome has disclosed new possibilities for the study of bio-cultural evolutionary processes. In this communication, I present a study on the relationships between social structure and genetic variation in sub-Saharan Africa based on published and unpublished mitochondrial and Y-chromosomal data relative to a total of 40 populations. The results obtained reveal a striking difference in the genetic structure of farming (Bantu speakers) and hunting-gathering (Pygmies and Bushmen) populations. To explain this difference, I propose that asymmetric gene flow, polyginy and patrilocality, and hence the socio-cultural factors underlying them, have had an important role in determining and differentiating the genetic structure of sub-Saharan populations. The limits and the implications of this study are discussed.     

Genetic structure and dispersal of Macrobrachium australiense (Decapoda: Palaemonidae) in western Queensland, Australia

SUMMARY 1. The freshwater prawn, Macrobrachium australiense (Decapoda: Palaemonidae), is an abundant species throughout the rivers of western Queensland, and it is thought to have effective dispersal capabilities. Given the very low topographical relief of the western Queensland landscape and the vast nature of episodic flooding, it was predicted that widespread dispersal in this species would occur within, and possibly between the four major catchments of the region: the Darling, Bulloo, Cooper and Diamantina. We analysed eight polymorphic allozyme systems and a fragment of the cytochrome c oxidase subunit 1 (COI) mitochondrial DNA (mtDNA) gene to determine the extent of recent and historical patterns of dispersal at nested spatial scales, within and between catchments.
2. Large and significant levels of allozyme and mtDNA differentiation were revealed among all catchments, indicating that dispersal of M. australiense does not occur across catchment boundaries, although this species is reportedly capable of overland movement. In contrast, no significant patterns of genetic differentiation were resolved between major subcatchments of the Darling and Cooper, or between sites within these subcatchments, indicating that populations of M. australiense are panmictic within catchments.
3. The MtDNA data resolved two divergent and reciprocally monophyletic clades, with the first representing the Darling catchment, and the second corresponding to the Bulloo, Cooper and Diamantina catchments. We postulate that extreme variation in historical climatic patterns and palaeohydrologic conditions played an important role in shaping the population structure of M. australiense throughout western Queensland during the Quaternary.     

Effect of water temperature on the mass emergence of the mayfly, Ephoron shigae, in a Japanese river (Ephemeroptera: Polymitarcyidae)

1. Winged stages of Ephoron shigae were collected every day during their emergence period during a 6-year period from 1989 to 1994, by net sweeps or a light trap along the Asahi-gawa River in western Japan. Emergence occurred mainly in September. 2. Coefficients of determination (r²) were calculated for the regression of the mean date of emergence against cumulative degree-days during various periods from late March to early September. 3. In both sexes the highest values of r² were obtained for regression with degree-days from late June to late August. This indicates that thermal conditions during the late instars affected the emergence timing most strongly. The timing of emergence can be well predicted from the cumulative degree-days during summer.     

Genetic population structure of red snappers (Lutjanus malabaricus Bloch & Schneider, 1801 and Lutjanus erythropterus Bloch, 1790) in central and eastern Indonesia and northern Australia

The genetic population structure of red snapper Lutjanus malabaricus and Lutjanus erythropterus in eastern Indonesia and northern Australia was investigated by allozyme electrophoresis and sequence variation in the control region of mtDNA. Samples were collected from eight sites in Indonesia and four sites in northern Australia for both species. A total of 13 allozyme loci were scored. More variable loci were observed in L. malabaricus than in L. erythropterus . Sequence variation in the control region (left domain) of the mitochondrial genome was assessed by RFLP and direct sequencing. MtDNA haplotype diversity was high ( L. erythropterus , 0·95 and L. malabaricus , 0·97), as was intraspecific sequence divergence, ( L. erythropterus , 0·0–12·5% and L. malabaricus , 0·0–9·5%). The pattern of mtDNA haplotype frequencies grouped both species into two broad fisheries stocks with a genetic boundary either between Kupang and Sape ( L. malabaricus ) or between Kupang and Australian Timor Sea ( L. erythropertus ). The allozyme analyses revealed similar boundaries for L. erythropterus . Seven allozymes stocks compared to two mtDNA stocks of L. malabaricus including Ambon, which was not sampled with mtDNA, however, were reported. Possible reasons for differences in discrimination between the methods include: i) increased power of multiple allozyme loci over the single mtDNA locus, ii) insufficient gene sampling in the mtDNA control region and iii) relative evolutionary dynamics of nuclear (allozyme loci) and mitochondrial DNA in these taxa. Allozyme and haplotype data did not distinguish separate stocks among the four Australian locations nor the central Indonesian (Bali and Sape locations) for both L. malabaricus and L. erythropterus.     

Isolation and characterization of 10 highly polymorphic di- and trinucleotide microsatellite markers in the mayfly Ameletus inopinatus (Ephemeroptera: Siphlonuridae)

We describe the isolation of ten polymorphic microsatellite loci from the mayfly Ameletus inopinatus. Loci had di‐ or trinucleotide repeat motifs and were highly variable with three to 17 alleles (mean = 7.15). Observed heterozygosity ranged from 0.143 to 0.905. One locus (Ami_202) showed significant deviation from Hardy–Weinberg equilibrium in one population, but no evidence for null alleles. One locus (Ami_73) was significantly linked with three other loci. The remaining nine loci should prove highly informative for population genetic studies.     

Genetic diversity, population structure, and conservation of Sophora moorcroftiana (Fabaceae), a shrub endemic to the Tibetan Plateau

Sophora moorcroftiana is a perennial leguminous low shrub endemic to the middle reaches of Yarlung Zangbo River in Tibet. It is an important species to fix sand dunes and to avoid the formation of shifting sands; therefore, its progressive over-exploitation may enhance land desertification. The levels and distribution of genetic variability of this species were evaluated from 10 natural populations at 24 loci encoding 13 enzymes, using allozyme analysis by starch gel electrophoresis. Data obtained revealed moderate levels of genetic variation within populations (Pp=27.5%, Ap=1.5, Hep=0.122) and a considerable divergence among populations (FST=0.199). Significant positive correlations (r2=0.49, p<0.05; r2=0.46, p<0.05) were found between elevation and both mean number of alleles per locus (A) and gene diversity (He) in the studied populations of S. moorcroftiana. Lower genetic diversity in lower elevation populations might be due to the negative effects of human pressures and habitat fragmentation, to adaptation to high altitudes as a consequence of a peripatric speciation process, or to directional gene flow along the river basin from the source populations located in the west at higher altitudes. The evaluation of the degree of threat has led to the inclusion of this species in the category of EN ("endangered"), and conservation strategies for this endemic species are discussed on the basis of these findings.     

Early embryonic development of the mayfly Ephemera japonica McLachlan (Insecta: Ephemeroptera,Ephemeridae)

In the newly laid egg of the mayfly Ephemera japonica, an egg nucleus (oocyte nucleus) at metaphase of the first maturation division is in the polar plasm at the mid-ventral side of the egg, and a male pronucleus lies in the periplasm beneath a micropyle situated just opposite the polar plasm or at the mid-dorsal side of egg. The maturation divisions are typical. An extensive and circuitous migration of the male pronucleus is involved in the fertilization process: it first moves anteriad in the periplasm from beneath the micropyle to the anterior pole of the egg and then turns posteriad in the yolk along the egg's long axis to the site of syngamy, near the center of the egg. Cleavage is superficial. The successive eight cleavages, of which the first five are synchronized, result in the formation of the blastoderm, and about ten primary yolk cells remain behind in the yolk. Even in the newly formed blastoderm, the thick embryonic posterior half and the thin extraembryonic anterior half areas are distinguished: the former cells are concentrated at the posterior pole of the egg to form the germ disc, and the latter cells become more flattened, forming serosa. Time-lapse VTR observations reveal a yolk stream that is in accord with the migration of the male pronucleus in time and direction. The yolk stream is also generated in activated unfertilized eggs, and it is probable that the migration of the male pronucleus in association with the fertilization may be directed by the yolk stream. J. Morphol. 238:327–335, 1998. © 1998 Wiley-Liss, Inc.     

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.     

Predation on mayfly nymph, Baetis rhodani, by native and introduced Gammarus: direct effects and the facilitation of predation by salmonids

1. In a series of laboratory experiments, we assessed the predatory nature of the native Irish amphipod, Gammarus duebeni celticus , and the introduced G. pulex , towards the mayfly nymph Baetis rhodani . We also investigated alterations in microhabitat use and drift behaviour of B . rhodani in the presence of Gammarus , and indirect predatory interactions with juvenile Atlantic salmon, Salmo salar .
2. In trials with single predators and prey, B. rhodani survival was significantly lower when Gammarus were free to interact with nymphs as than when Gammarus were isolated from them. The invader G. pulex reduced the survival of B. rhodani more rapidly than did the native G. d. celticus . Both Gammarus spp. were active predators.
3. In `patch' experiments, B. rhodani survival was significantly lower both when G. pulex and G. d. celticus were present, although the effect of the two Gammarus species did not differ. Again, active predation of nymphs by Gammarus was observed. Significantly more nymphs occurred on the top and sides of a tile, and per capita drifts were significantly higher, when Gammarus were present. Baetis rhodani per capita drift was also significantly higher in the presence of the introduced G. pulex than with the native G. d. celticus .
4.  Gammarus facilitated predation by salmon parr of B. rhodani by significantly increasing fish–nymph encounters on exposed gravel and in the drift. There were no differential effects of the two Gammarus spp. on fish – B . rhodani encounters or consumption.
5. We conclude that Gammarus as a predator can have lethal, nonlethal, direct and indirect effects in freshwaters. We stress the need for recognition of this predatory role when assigning Gammarus spp. to a `Functional Feeding Group'.     

Long‐term monitoring of the life history of a riverine mayfly Ephemera orientalis (Ephemeroptera: Ephemeridae) in the Han River in Seoul,Korea

Climatic change often affects life history aspects of aquatic insects. Long‐term monitoring was conducted to understand the life history variability of the riverine mayfly Ephemera orientalis in the Han river, where habitat is largely disturbed by summer floods. Water level of the study site, Godeok‐dong area in Seoul, is regulated by the Paldang Dam located approximately 14 km above the study site. E. orientalis often emerged in large numbers around the study area. Larvae were sampled monthly from April 2006 to July 2010 using a Surber sampler (50 × 50 cm, mesh 0.25 mm, two replicates). Additional qualitative sampling was conducted using a hand net. As a result, based on the larval body size distribution, E. orientalis had a univoltine life cycle with two distinct slow growing cohort groups: the S1group emerged during May–June and the S2 group emerged during August–September. Our previous study conducted in Gapyeong stream in Gyeonggi‐do showed that E. orientalis normally had three cohort groups, S‐1 and S‐2 groups and the F‐group (a fast growing cohort developed during May–August). However, the Han River population of E. orientalis lacked such a fast growing cohort. It is most probable that habitat disturbances caused by high water level during the rainy season negatively affected the development of the fast growing cohort (F‐group) in the Han River area. The relationships between monthly E. orientalis population data and water level in the Han River are presented.