Genome size evolution and polyploidy in the Daphnia pulex complex (Cladocera: Daphniidae)

Genome size was estimated in 49 clones of the Daphnia pulex complex from temperate and subarctic locations using flow cytometry and microsatellite DNA analyses. Significant genome size differences were found in diploid species belonging to the two genetically distinct groups (the pulicaria and the tenebrosa groups), with clones from the tenebrosa group having genome sizes 22% larger than those in the pulicaria group. Combined flow cytometry and microsatellite DNA analyses revealed that nearly all polyploid clones in the D. pulex complex are triploid and not tetraploid, as was previously suggested. Sequencing analyses of the ND5 gene to position clones in their respective clades within the D. pulex complex have uncovered three triploid clones of Daphnia middendorffiana with a D. pulex maternal parent. This result was unexpected because Daphnia pulicaria has always been identified as the maternal parent of these hybrid polyploid clones. Triploid clones likely owe their origins to interactions between sexual and asexual populations. Further interactions in the tenebrosa group have generated tetraploid clones but these events have been rare.  © 2009 The Linnean Society of London, Biological Journal of the Linnean Society , 2009, 97 , 68–79.     

Carapace epithelia are rich in large filamentous actin bundles in Daphnia magna,Daphnia pulex,and Sida crystallina (Crustacea: Cladocera)

Cladocerans (water fleas) are planktonic crustaceans that typically have a bivalved carapace. Each valve of the carapace consists of two cuticle‐secreting epithelial layers that are separated by a hemolymphatic chamber and joined by pillar structures. Ultrastructural analyses in several species of Cladocera have shown that the carapace epithelia and pillars contain filamentous structures of unknown composition. In the present study we used a fluorescent phalloidin conjugate to show that the carapaces of three cladocerans, Daphnia magna, D. pulex, and Sida crystallina, are rich in large bundles of filamentous actin (F‐actin). In D. magna we employed confocal microscopy and orthogonal views of three‐dimensional reconstructions to show that these bundles extend radially from foci in the pillars towards the integument surfaces, and their structure is consistent with that of contractile stress fibers. Using a fluorescent lipophilic stain, DiOC₆(3), we show that the F‐actin bundles are distributed in membrane‐rich regions within the carapace epithelia, and that, in the superficial epithelium, these may be large membrane‐bound organelles. In D. magna, the F‐actin bundles are present in embryonic, juvenile instar, and adult, developmental stages, and through development the bundles become larger, contain more F‐actin, and become more widely spaced. We present an alignment of the deduced amino acid sequences of six putative D. pulex actin genes, and discuss the implications that their respective sequences have on the likelihood of their inclusion into the F‐actin bundles of the carapace. Our identification of these large F‐actin bundles within the pillars of three cladocerans provides new insight into the role these structures play in influencing carapace dynamics within this order.     

Daphnia curvirostris Eylmann in Japanese high mountain waters (Cladocera: Daphniidae)

A species of Daphnia, Daphnia curvirostris Eylmann, found in high mountain lakes and ponds in central Japan is described. Although there were some differences in the shape of the male rostrum and the chromosome number between European populations as described by Johnson (1952) and Trentini (1980), and Japanese ones collected from high mountain waters, Japanese specimens had many characteristics similar to the taxon D. curvirostris of Europe.     

Fish kairomone regulation of internal swarm structure in Daphnia pulex (Cladocera: Crustacea)

In this laboratory experiment, swarms of D. pulex were artificially created by placing thirty individuals in small chambers containing 130 ml of water. The swimming behaviour of the animals was measured using a video camera and a programme to digitize observations. The D. pulex swimming in water with fish kairomones had a more uniform swimming speed compared with animals in control water. If aggregated prey individuals benefit from a reduced predation risk, and this risk is further reduced by uniformity of swarm members, the uniformity of swimming speed can be interpreted as an behavioural adjustment to minimize the vulnerability to predation. This revised version was published online in July 2006 with corrections to the Cover Date.     

A comparison of populations from the Daphnia similis group (Cladocera: Daphniidae)

European populations of Daphnia similis Claus have been compared with populations from tropical Asia. Daphnia similoides n. sp. from tropical Asia was described as the sibling species of D. similis. The female has characteristic neonate, postabdomen, head, and ephippium. The male has characteristic rostrum, antennules, and the distalmost part of the postabdomen. The D. similis group was compared with D. carinata s. str. Australian populations.     

The fine structure of the digestive system of Daphnia pulex (Crustacea: Cladocera).

The alimentary canal of Daphnia pulex consists of a tube-shaped foregut, a midgut (mesenteron) with an anterior pair of small diverticula, and a short hindgut. The foregut and hindgut are structurally similar. Each is formed by a low cuboidal epithelium 5 mum tall and lined with a chitinous intima. The midgut wall consists of a simple epithelium resting on a thick beaded basal lamina which is surrounded by a spiraling muscularis. Anteriorly the midgut cells are columnar in shape being 30 mum in height each having a basal nucleus, anteriorly concentrated mitochondria and in apical border of long thin microvilli. Posteriorly the midgut cells become progressively shorter so that in the posteriormost region of the midgut the cells are 5 mum tall and cuboidal in shape. The microvilli concomitantly become shorter and thicker. All mesenteron cells contain the usual cytoplasmic organelles. The paired digestive diverticula are simple evaginations of the midgut. The wall of each consists of a simple epithelium of cuboidal cells 25 mum in height, each with a brushed border of long thin microvilli. Enzyme secretion appears to be holocrine in mode and not confined to any one region of the mesenteron though definitely polarized anteriorly. The thin gut muscularis encircles the entire length of the midgut and caeca. Thick and thin filaments appear to be in a 6:1 ratio.     

Morphology and age variability of Daphnia galeata Sars (Cladocera: Daphniidae) in two adjacent water bodies of the Kola Peninsula

The morphology and age variability of Daphnia galeata Sars (Cladocera: Daphniidae) in two adjacent water bodies of the Kola Peninsula are studied. The reliability of characters usually used for diagnosis of adult individuals of the given species was verified, and their formation in the postembryonic period was traced. If the identification of D. galeata from Lake Bolshoi Vudvyar caused no doubts, the population from Lake Malyi Vudvyar was atypical with respect to several morphological characters. We concluded that plastic characters, such as body shape, as well as the shape of the dorsal crest and helmet on the head are unreliable for distinguishing closely related species of water fleas. At the same time, some small-size characters, for instance, the relative depth of the medial crest on the posterior side of the head, allow us to diagnose this species with confidence.     

Morphological Radiation with Reference to the Carapace Valves of the Anomopoda (Crustacea: Cladocera)

Diversity of the carapace valve structure in the Anomopoda (Crustacea: Branchiopoda) is described. We found that transformations of shape, modifications of the surface and valve margins, and thickening or thinning of the valves, take place independently in different families, sometimes resulting in a convergent similarity. The limits of morphological diversity in these features determine the range of adaptive radiation in general form of the group. (© 2009 WILEY‐VCH Verlag GmbH & Co. KGaA, Weinheim)     

Other environmental factors affect the size of crests developed by Daphnia carinata (Cladocera: Daphniidae) in response to notonectid predators

SUMMARY. 1. Crest size and body size was measured in Daphnia carinata King sampled from field enclosures in which environmental conditions and notonectid predator ( Anisops Hyperion Kirkaldy) levels had been manipulated, and from control treatments which had predator levels and other environmental factors equivalent to those in the natural environment.
2. Individuals of D. carinata developed larger crests in treatments with predators than in treatments without predators.
3. The environmentally manipulated treatment was considered to have lower levels of food but was also likely to have undergone other changes in physicochemistry and resource variability that may have affected crest size. The effect of the manipulation cannot be confirmed, therefore, as the result of variation in food levels. Animals in the treatments considered to have lower amounts of food had smaller crests than in the control treatment irrespective of whether predators were present. The effects of environmental manipulation and predators were additive.
4. Enclosures without predators, but in which water from the natural, predator-rich, environment was exchanged through the enclosure walls, showed no evidence of a chemical induction effect. Chemical induction effects may have been countered by a reduced food supply resulting from the higher D. carinata population densities that developed in these enclosures.
5. This work provides a second example of an alteration of the morphological response of daphnid prey to predators by environmental factors, probably food, and suggests this phenomenon may be general in cladoceran species displaying predator-induced changes in morphology.     

Molecular genetic analysis of Daphnia in the English Lake District: species identity, hybridisation and resting egg banks

• 1 Taxonomic issues in many Daphnia species complexes are often confused by a high degree of phenotypic plasticity and by interspecific hybridisation. Here, we employ molecular genetic tools to confirm the species composition and incidence of hybridisation in extant and resting egg populations of Daphnia from Windermere and Esthwaite Water in the English Lake District.
• 2 A combination of species‐diagnostic allozymes and mtDNA, confirms that contemporary populations are dominated by a single species, D. galeata.
• 3 We present the first account of genetic characterisation of dated ‘resting’ eggs using microsatellites and mtDNA, employing PCR‐based DNA recovery, thus providing a temporal dimension to taxonomic patterns. Thirty years ago, two species were present in Esthwaite, D. galeata and D. hyalina, but Windermere populations were dominated by D. galeata only.
• 4 The use of PCR‐based mtDNA RFLP analysis as a species‐diagnostic tool, and microsatellites to monitor clonal diversity, provide a valuable approach for long‐term studies, especially in populations free from the complicating effect of frequent hybridisation. The detailed limnological records available for many large lakes, and associated changes in land‐use, pollutants and climate, combined with long‐term ephippial molecular genetic data, provide opportunities for exploring natural and anthropogenic impacts on genetic and community structure.

     

Wind-borne short-range egg dispersal in anostracans (Crustacea: Branchiopoda)

Evidence is provided for the first time that at least part of the egg bank of the southern African fairy shrimp Branchipodopsis wolfi is dispersed over short distances by wind. A total of 423 sticky surfaces were mounted around and between individual basins at three rock pool sites, to trap any dispersing eggs during a 3 day period. Eight viable eggs were found in different egg traps (1.9% of the total), seven of which were located at one site with shallow basins. Given the proximity of vast egg banks to the egg traps, the low observed dispersal rate cautions against overestimating the importance of wind dispersal for population genetic processes. By generating egg banks to hedge against drought catastrophes, and producing egg types with different potential dispersibility, B. wolfi is a strong bet-hedger which has established a means of escaping temporal and spatial stress. These life history functions enable populations to persist in small desert rock-pools, which are the most extreme of temporary habitats.     

Sexual reproduction in Daphnia pulex (Crustacea: Cladocera): observations on male mating behaviour and avoidance of inbreeding

1.  Mating behaviour in Daphnia appears to rely on random contact between males and sexual females rather than diffusible pheromones. Males may be able to discriminate sexually receptive females from females in other developmental stages and increase their mating efficiency. Males may also use chemical signals to avoid mating with females from the same clone and avoid the severe inbreeding depression that has been documented for intraclonal mating. The present study used experiments to test for the avoidance of intraclonal mating and assess male mating efficiency in D. pulex .
2.  Three clones were examined for the avoidance of intraclonal mating by providing males with an opportunity to mate with females of the same or two different clones. The proportion of intraclonal matings did not differ from the proportion of interclonal matings, suggesting that D. pulex males do not use kin discrimination to avoid mating with females from the same clone.
3.  The proportion of mated females decreased with increasing numbers and density of sexual females when exposed to a single male. This observation suggests that a male spends more time pursuing and copulating with sexually receptive females than non-receptive females and there is insufficient time to mate with all sexual females. The decrease in proportion of females mated could also be the result of sperm depletion in the male. Sperm depletion is unlikely to occur in nature because sexually receptive females are much rarer than in the experimental conditions.     

Desiccation and egg viability of the ragwort flea beetle,Longitarsus flavicornis (Coleoptera: Chrysomelidae)

A study was conducted to investigate the effect of desiccation on the survival of eggs of Longitarsus flavicornis. Eclosion of L. flavicornis eggs in laboratory trials decreased with increasing desiccation time between 0 days (93% hatching) and 42 days (no egg hatching) at 50±2% relative humidity and 23±2°C. Probit analysis indicated that 25, 50 and 99% mortality of L. flavicornis eggs occurred after 5.7, 9.3 and 50.4 days desiccation, respectively. Egg development varied between a minimum of 8 days at 7 days desiccation to a maximum of 15 days at 28 days desiccation. Hatching span did not differ between treatments with all eggs hatching within 12 days of each other. A relative humidity of 88–100% was measured under ragwort rosettes in non-drought field conditions. This would be expected to facilitate successful egg eclosion. However, the occurrence of summer drought could be detrimental to egg survival.     

Lifetime egg production and egg mortality in the damselflyPyrrhosoma nymphula (Sulzer) (Zygoptera: Coenagrionidae)

Artificial oviposition sites were used to estimate egg deposition rates in the field. Females laid an average of 10.76 eggs/minute with a mean duration of 22.81 minutes, giving an average clutch size of 245 eggs. Since one mating corresponded to one clutch of eggs, lifetime mating success was used as a measure of the number of clutches produced. Mean lifetime clutch production was 5.91 clutches per female, equating to 1447 eggs per female per lifetime. Eggs were hatched in the laboratory at temperatures comparable with those in the field. Hatching was highly synchronised and the overall hatching success was 75.1%. Causes of egg mortality in the laboratory were limited to infertility and unhatchability. Since no other sources of egg mortality could be found at the study site, this value was a good reflection of hatching success in the field. Lifetime egg production and hatching success were used to estimate the number of viable offspring produced per female, giving a higher order estimate of reproductive success than has previously been published for a zygopteran.     

Diurnal vertical migration of Daphnia hyalina Leydig, 1860 (Crustacea: Cladocera) in Lake Bled (Slovenia) in relation to temperature and predation

Diurnal vertical migration (DVM) of Daphnia hyalina in Lake Bled was most intense during summer stratification. The extent of DVM varied with the size of the animal and its reproductive state. Migration distances were shortest in immature specimens and longest in ovigerous females. During daytime, ovigerous females stayed deeper in the water column than females without ova or immatures. The daytime temperature of water at the median depth of the ovigerous females did not exceed 10 °C, even in the warmest season. At night they migrated upward to an environment which was warmer by as much as 9 °C.Laboratory observations indicate that specimen's size and water temperature determine the velocity of passive sinking, such that morning descent of the different groups of Daphnia can be explained by passive sinking alone.Our hypothesis is that the distribution of different groups of D. hyalina in Lake Bled is influenced by two types of predators: fish (Perca fluviatilis L. and Rutilus rutilus (L.)) and larvae of Chaoborus flavicans (Meig.), the latter appearing in the epilimnion during the night. Fish predation has a key-role at the beginning of thermal stratification. Supposing that in spring the gene pool of Daphnia consists of a mix of different genotypes, distributed at different depths during the day, fish predation combined with a presence of fish chemicals favored genotypes with a lower day-depth during the spring/summer period.     

The distribution and variation of Daphnia lumholtzi (Crustacea: Cladocera) in relation to fish predation in Lake Albert, East Africa

In Lake Albert Daphnia lumholtzi is found in two forms. One has a pointed anterior prolongation, or helmet, on the head. The other has a shorter rounded head, and was originally described as a separate species, D. monacha . The latter form dominates the zooplankton in the middle of the lake where planktivorous fish are rare or absent. The helmeted form becomes commoner near the margins of the lake and reaches its greatest abundance in Ndaiga Lagoon, where planktivorousfish are common. The possession of a helmet is associated with a reduction in the size of the carapace compared to the round headed form. The carapace with its contained eggs is the most conspicuous part of a cladoceran, so that the helmeted forms are at an advantage in the presence of planktivorous fish which locate their prey by sight. The mid-lake monacha forms are larger than specimens of the same form in Ndaiga Lagoon, where it is shown that Alestes baremose feeds selectively on the larger specimens of the monacha form.
The helmeted form produces more, but smaller eggs than the monacha form. The total brood volume (= mean egg volume x mean number eggs per female) is greatest in the midlake monacha forms. The selective advantages of variations in egg size and the possession of a helmet are discussed. It is concluded that the data from Lake Albert support the hypothesis of Brooks (1965) concerning the adaptive significance of helmet development in Daphnia .