The systematics of Australian Daphnia (Cladocera: Daphniidae). Multivariate morphometrics

Forty nine characters were measured in each of 76 male and 200 female Daphnia including specimens from all eleven taxa described from the genus in Australia, except D. jollyi. Separate multivariate analyses (cluster, principal components and discriminant) of males and females each revealed only five morphological groups in the genus corresponding to the species D. occidentalis, D. lumholtzi, D. cephalata, D. nivalis, and a conglomerate of all other taxa described from the D. carinata complex, D. carinata (sensu lato). Interpretation of the male results was straightforward, but that for females was not because of the occurrence of an environmentally induced seasonal change in morphology (cyclomorphosis) in females. Log transformations of the female data successfully isolated the effects of cyclomorphosis. The seasonal morphs formed two groups which were further differentiated into taxonomic groups.     

Inducible defences in Daphnia: responses to two closely related predator species

The predator-induced responses of two species of Australian Daphnia, with contrasting distributions and life history patterns, to the kairomones of two species of Anisops predators, were measured. Daphnia longicephala produced a large crest and attained a larger size when exposed to both predators. D. carinata sl matured earlier than D. longicephala and did not produce a crest. Surprisingly, kairomones of both predators inhibited the production of ephippia in D. carinata sl. Anisops stali, the larger of the two predator species, induced a significantly larger crest size in D. longicephala, and larger brood size in both species compared with the smaller A. gratus, indicating a quantitative but not qualitative effect of predator species on inducible defences. Received: 18 August 1999 / Accepted: 14 April 2000     

Characteristics of six cladocerans in relation to ecotoxicity testing

Investigation was made to compare some biological characteristics relevant to ecotoxicity testing among six cladoceran species, including Daphnia magna, Daphnia carinata, Daphnia pulex, Ceriodaphnia quadrangular, Bosmina longirostris, and Simocephalus vetulus. The results show that D. carinata had advantages over other cladoceran species for being used as a test organism, particularly for ecotoxicological assessment of aquatic environments in tropical and subtropical areas. D. carinata had similar body size and total number of offspring per female to D. magna. However, D. carinata was more sensitive to the reference toxicant and had much shorter reproduction cycle than D. magna. D. carinata had similarity to D. pulex, C. quadrangular, S. vetulus and B. longirostris in terms of sensitivity to the reference toxicant and length of reproduction cycle. However, D. carinata was much larger in size and produced much more offspring per female than any of D. pulex, C. quadrangular, S. vetulus and B. longirostris. Among the investigated cladocerans, only the neonates (2- and 4-day-old) of D. carinata exhibited phototaxis that was sufficiently remarkable. The low among-generation variation in phototaxis index (I_p) of D. carinata and the close relationship between I_p and the concentration of the ISO standard toxicant (K₂Cr₂O₇) appears to suggest that I_p can be used as an excellent test endpoint for ecotoxicity testing.     

The systematics of Australian Daphnia (Cladocera: Daphniidae). Electrophoretic analyses of the Daphnia carinata complex

Genetic variation at eleven allozyme loci was scored in approximately 1600 individual Daphnia. The samples included representatives of all nine taxa described from the D. carinata complex in Australia. Principal coordinates and cluster analyses revealed only three groups of genotypes within the complex, corresponding to the taxa D. cephalata, D. nivalis, and a conglomerate of all other taxa described from the complex, D. carinata (sensu lato). These results are consistent with recent multivariate morphological analyses of the complex.     

The response of European Daphnia magna Straus and Australian Daphnia carinata King to changes in geomagnetic field

This study investigates the effects of lifelong exposure to reversed geomagnetic and zero geomagnetic fields (the latter means absence of geomagnetic field) on the life history of Daphnia carinata King from Australia and Daphnia magna Straus from Europe. Considerable deviation in the geomagnetic field from the usual strength, leads to a decrease in daphnia size and life span. Reduced brood sizes and increased body length of neonates are observed in D. magna exposed to unusual magnetic background. The most apparent effects are induced by zero geomagnetic field in both species of Daphnia. A delay in the first reproduction in zero geomagnetic field is observed only in D. magna. No adaptive maternal effects to reversed geomagnetic field are found in a line of D. magna maintained in these magnetic conditions for eight generations. Integrally, the responses of D. magna to unusual geomagnetic conditions are more extensive than that in D. carinata. We suggest that the mechanism of the effects of geomagnetic field reversal on Daphnia may be related to differences in the pattern of distribution of the particles that have a magnetic moment, or to moving charged organic molecules owing to a change in combined outcome and orientation of the geomagnetic field and Earth's gravitational field. The possibility of modulation of self-oscillating processes with changes in geomagnetic field is also discussed.     

A study on the effects of food quantity and quality on glutathione S-transferase (GST) activity and growth rate parameters of <Emphasis Type="Italic">Daphnia</Emphasis><Emphasis Type="Italic">carinata</Emphasis> varying in age

The adverse influences of insufficient food and toxins on Daphnia carinata’s body growth, reproduction and tolerance were investigated in the laboratory. Different concentrations of Scendesmus obliquus and a mixture of S. obliquus and microcystin (MC)-containing Microcystis aeruginosa PCC7820 were used to feed D. carinata. Glutathione S-transferase (GST) activity towards five chemical compounds (substrates) was measured and used as an indicator of their tolerance. Body growth rate and clutch size of D. carinata decreased with declined concentration or decreased proportion of S. obliquus in the diet. GST activity decreased with ageing in D. carinata. However, GST activity to several chemical compounds increased when food quantity or food quality decreased. Adult D. carinata had a lower GST activity towards p-nitrophenenzyl chloride (PNBC) than juveniles and exhibited a sharp decline (P < 0.001) in GST activity towards PNBC as the animals aged. It is suggested that the age-specific decrease in GST activity is a possible mechanism for the high morality of adult D. carinata in the summer of eutrophic lakes.     

Polymorphism and Food Limitation in Three Daphnia carinata Populations

The annual cycles of Daphnia carinata in three fish-free ponds in southern New Zealand (Quarry, Taieri and Raupo Ponds) are described. Although breeding was continuous, the populations were frequently dominated by distinct cohorts and there was a sexual phase in spring. Growth rates, particularly in Quarry Pond, were often slow, and clutch sizes generally small. Egg and neonate sizes changed seasonally, but changes were not closely related to clutch sizes and body lengths. In Taieri Pond and Raupo Pond some D. carinata grew to 6.14 mm and carried clutches of up to 154 eggs. The potentially high fecundity associated with large size makes D. carinata a good coloniser, but our studies showed that it also survives well in poor food conditions by growing slowly and producing only 1–2 eggs per clutch. D. carinata showed marked seasonal polymorphism in tailspine length and development of head crests. In winter, crests were absent and tailspines were short and varied little with body length. Crests were first apparent, and tailspines began to elongate, in early spring, before the first appearance of a notonectid, Anisops wakefieldi. By midsummer, tailspines had increased markedly with body length but the slope of the relationship declined in autumn. Laboratory trials showed that both tailspine length and crest development increased in the presence of Anisops. These increases may be due to water-soluble factor(s) produced by Anisops.     

Seasonal variation in life history traits in two Drosophila species

Seasonal environmental heterogeneity is cyclic, persistent and geographically widespread. In species that reproduce multiple times annually, environmental changes across seasonal time may create different selection regimes that may shape the population ecology and life history adaptation in these species. Here, we investigate how two closely related species of Drosophila in a temperate orchard respond to environmental changes across seasonal time. Natural populations of Drosophila melanogaster and Drosophila simulans were sampled at four timepoints from June through November to assess seasonal change in fundamental aspects of population dynamics as well as life history traits. D. melanogaster exhibit pronounced change across seasonal time: early in the season, the population is inferred to be uniformly young and potentially represents the early generation following overwintering survivorship. D. melanogaster isofemale lines derived from the early population and reared in a common garden are characterized by high tolerance to a variety of stressors as well as a fast rate of development in the laboratory environment that declines across seasonal time. In contrast, wild D. simulans populations were inferred to be consistently heterogeneous in age distribution across seasonal collections; only starvation tolerance changed predictably over seasonal time in a parallel manner as in D. melanogaster. These results suggest fundamental differences in population and evolutionary dynamics between these two taxa associated with seasonal heterogeneity in environmental parameters and associated selection pressures.     

The costs of crest induction for Daphnia carinata

The effects of notonectid-induced crests on growth and reproduction, and resource allocation to crest construction, moult losses and eggs of Daphnia carinata were measured. An attempt was made to elucidate the mechanisms of physiological costs of crest induction for this species. The crested morph of d. carinata reached a significantly larger size than the uncrested form. Reproductive output was similar in early instars, but the crested morph produced more eggs in latter broods. Instar duration was longer for the crested morph and age at first reproduction was delayed. Survival was also lower in this form. Crest construction required significant resources (equivalent to 60 eggs over a life time) but evidence is presented that these resources were obtained primarily by re-allocation of available material rather than collection of extra resources. The crested morph allocated significantly more resources to moulting than its uncrested counter-parts. The uncrested morph produced large eggs in early instars and progressively smaller ones in later instars. The crested morph produced only small eggs. The hypothesis is presented that the crest-induction strategy of D. carinata involves at least two separate sets of responses, each with its own costs and trade-offs. The first response is production of the crest. The cost of crest production is an increased cost of moulting. D. carinata off sets this cost by increasing instar duration and thus age at reproduction. The second response is increased size. D. carinata achieves this by reducing the fraction of available resources allocated to reproduction. The cost is lower reproductive output.     

Effect of the epizoic rotifer Brachionus rubens on the population growth of three cladoceran species

Using population densities and growth rates as criteria, we studied interactions between the epizoic rotifer Brachionus rubens and each of three cladoceran species differing in size and reproductive rates — Daphnia carinata, Moina macrocopa and Ceriodaphnia rigaudi. In all mixed — species experiments, B. rubens existed in both the epizoic mode, attached to the cladoceran host, and in the free-swimming mode. Rotifer population growth rates were significantly depressed in the presence of M. macrocopa, presumably as a consequence of exploitative and interference competition. The largest cladoceran, D. carinata probably did not suppress B. rubens, because the epizoic component of the rotifer population escaped from the deleterious effects of mechanical interference. Peak population numbers and initial population growth rates reached by all three cladocerans were lower in the presence of B. rubens, probably because of the adverse effects of the epizoic infestation, which was maximal on D. carinata and least on C. rigaudi. In mixed-species cultures of D. carinata and M. macrocopa, the presence of B. rubens helped D. carinata coexist with M. macrocopa, which otherwise would have suppressed the Daphnia.     

Elemental composition of two coexisting Daphnia species during the seasonal course of population development in Lake Constance

Summary In 1986 and 1987 the seasonal course of population density, life history traits and biomass composition (CHN-elemental and ash) of two coexisting Daphnia species from L. Constance (Überlinger See) were studied. The two daphnids are morphologically similar, but demonstrate different seasonal variation of behaviour (i.e. diurnal vertical migration). With the exception of nitrogen content, the mean biomass composition of adults and that of eggs were significantly different: the dry biomass of eggs (adults) contained on average 55.6 (46.1) % C, 7.9 (6.5) % H, 9.7 (9.7) % N, and 10.0 (23.8) % ash. These values were similar to those of other daphnids; it may be concluded that the average daphnid composition is not species specific, but is determined mainly by the position in the life cycle and by the nutritional state of the animal. A distinct seasonal variation in the elemental composition of the biomass of adult daphnids was observed. It related significantly to the concentration of food, as indicated by the epilimnetic chlorophyll-a concentrations and by Secchi depth. From the amplitude of seasonal variation of elemental composition and from the shape of relationships between the elemental composition of daphnids and their food, it can be concluded that in summer the non-migrating D. galeata lives under stronger food limitation than the migrating H. hyalina. In spring, there were no differences in the biomass composition of either species. In summer, the adult fcmales of D. galeata contained significantly less carbon, and hydrogen than those of D. hyalina. In most parameters of biomass (i.e. biomass composition of adults) and of life history (i.e. clutch size, mean adult weight), D. galeata displayed higher coefficients of variation. A tendency in D. galeata to have peaks of population development carlier in the season and in D. hyalina to have them towards autumn was observed. These contrasting responses of the populations development, the seasonal variability of the biomass composition seem to support the view of Geller (1986) who called D. galeata and D. hyalina exploitative and conservative strategists respectively.This research was supported by the Deutsche Forschungsge-meinschaft within the Sonderforschungsbereich Cycling of Matter in Lake Constance (SFB 248), and by scholarship to R.B. from the Deutscher Akademischer Austauschdienst (DAAD)     

Temperature and kairomone induced life history plasticity in coexisting Daphnia

We investigated the life history alterations of coexisting Daphnia species responding to environmental temperature and predator cues. In a laboratory experiment, we measured Daphnia life history plasticity under different predation risk and temperature treatments that simulate changing environmental conditions. Daphnia pulicaria abundance and size at first reproduction (SFR) declined, while ephippia (resting egg) formation increased at high temperatures. Daphnia mendotae abundance and clutch size increased with predation risk at high temperatures, but produced few ephippia. Thus, each species exhibited phenotypic plasticity, but responded in sharply different ways to the same environmental cues. In Glen Elder reservoir, Kansas USA, D. pulicaria dominance shifted to D. mendotae dominance as temperature and predation risk increased from March to June in both 1999 and 2000. Field estimates of life history shifts mirrored the laboratory experiment results, suggesting that similar phenotypic responses to seasonal cues contribute to seasonal Daphnia population trends. These results illustrate species-specific differences in life history plasticity among coexisting zooplankton taxa.     

The effect of photoperiod on life history and blood‐feeding activity in Aedes albopictus and Aedes aegypti (Diptera: Culicidae)

Several studies have examined how climatic variables such as temperature and precipitation may affect life history traits in mosquitoes that are important to disease transmission. Despite its importance as a seasonal cue in nature, studies investigating the influence of photoperiod on such traits are relatively few. This study aims to investigate how photoperiod alters life history traits, survival, and blood‐feeding activity in Aedes albopictus (Skuse) and Aedes aegypti (Linnaeus). We performed three experiments that tested the effects of day length on female survival, development time, adult size, fecundity, adult life span, and propensity to blood feed in Ae. albopictus and Ae. aegypti. Each experiment had three photoperiod treatments: 1) short‐day (10L:14D), 2) control (12L:12D), and 3) long‐day (14L:10D). Aedes albopictus adult females were consistently larger in size when reared in short‐day conditions. Aedes aegypti adult females from short‐day treatments lived longer and were more likely to take a blood meal compared to other treatments. We discuss how species‐specific responses may reflect alternative strategies evolved to increase survival during unfavorable conditions. We review the potential impacts of these responses on seasonal transmission patterns, such as potentially increasing vectorial capacity of Ae. aegypti during periods of shorter day lengths.     

Predictors of invasion success by Daphnia species: influence of food, temperature and species identity

Non-indigenous North American Daphnia ‘pulex’ has recently invaded lakes and reservoirs in South Island, New Zealand, that formerly contained only native Daphnia carinata. New Zealand is characterised by a wide range of freshwater ecosystems and low species diversity of planktonic crustaceans, particularly Cladocera. The potential success of a species to invade and establish in a new community is likely to be predicted more accurately when reproductive response norms and fitness of key resident species, as well as the non-indigenous species, have been established under a range of relevant environmental conditions. Based on the results of experiments to test aspects of reproduction and fitness of the invader, D. ‘pulex’, and D. carinata when grown together at a range of relevant temperatures (8–23 °C) and related photoperiods, I predict the species-specific potential of D. ‘pulex’ to be dispersed and colonise New Zealand lentic habitats, and the potential of D. carinata to persist with the invader in these habitats. Larger population densities of D. ‘pulex’ compared to D. carinata at higher temperatures and food level, and larger densities of D. carinata at low temperatures, imply a potential for both species to coexist in New Zealand lakes, facilitated by seasonal succession; increased water temperature and nutrient input associated with climate and land use changes appear likely to promote the wider establishment of D. ‘pulex’, with both negative and positive implications for the conservation and management of New Zealand’s freshwater ecosystems.     

Filtering rates of Daphnia carinata King (Crustacea: Cladocera) on the blue-green algae Microcystis aeruginosa Kütz. and Anabaena cylindrica (Lemm.)

Abstract Non-toxic strains (by mouse toxin assay test) of unicellular Microcystis aeruginosa and short filamentous Anabaena cylindrica were fed to Daphnia carinata in the laboratory at an algal volume concentration of 4.0 mm³ L^?1 at 24 ± 1°C. The filtering rates (FR, mL animal^?1 hr^?1) of D. carinata on M. aeruginosa and A. cylindrica increased with increasing body length (L, mm), and were expressed as a power curve: FR = 0.061L²⁰⁹. and FR = 0.232L^2.09, respectively. The potential for control of natural blue-green algal populations by D. carinata grazing is discussed briefly.     

Taxon-specific reaction norms to predator cues in a hybrid Daphnia complex

1. Previous studies have shown that interspecific hybridisation is common among taxa from the Daphnia galeata/hyalina/cucullata species complex. We investigated the influence of predator kairomones on the morphology and life histories of nine clones belonging to three taxa (pure D. galeata, F₁ hybrids between D. galeata and D. hyalina, and backcrossed D. hyalina) of this species complex. Predators exerting positive (fish) and negative (Chaoborus larvae) size‐selective predation were tested. 2. The most responsive traits were size at maturity and size of neonates. Despite large between‐clone variation, discriminant analysis revealed that the three taxa were distinct from each other in key life‐history traits. F₁ hybrids did not react in an intermediate way compared to the other taxa: the multivariate distances between F₁ hybrids and either taxon were larger than between pure D. galeata and backcrossed D. hyalina. 3. The average plasticity (calculated across all traits) was similar for all three taxa. With regard to the size at maturity and neonate body size, the strength of the response was a function of the intrinsic values of these traits expressed in the control. For example, for size at maturity, smaller individuals showed a significantly stronger reaction to Chaoborus kairomones than larger ones. 4. Finally, we monitored seasonal changes in body size, egg number and population density of pure D. galeata and F₁ hybrids in Greifensee (Switzerland). The two taxa experienced similar seasonal changes in body size but, on some sampling dates, they differed in mean egg number. The observed seasonal changes in Daphnia body size were consistent with what would be expected if the predator assemblage shifted from fish to Chaoborus over the course of the summer. The fluctuations in the frequencies of Daphnia taxa, however, were not related to seasonal variation in Daphnia body size. 5. Experimental data suggest that temporally heterogeneous predation regimes might be an important condition stabilising the co‐occurrence of Daphnia hybrids with parental taxa. Predation regimes, however, cannot solely explain dynamic changes in taxon frequency in Greifensee.     

PLEIOTROPY IN THE WILD: THE DORMANCY GENE DOG1 EXERTS CASCADING CONTROL ON LIFE CYCLES

In the wild, organismal life cycles occur within seasonal cycles, so shifts in the timing of developmental transitions can alter the seasonal environment experienced subsequently. Effects of genes that control the timing of prior developmental events can therefore be magnified in the wild because they determine seasonal conditions experienced by subsequent life stages, which can influence subsequent phenotypic expression. We examined such environmentally induced pleiotropy of developmental‐timing genes in a field experiment with Arabidopsis thaliana. When studied in the field under natural seasonal variation, an A. thaliana seed‐dormancy gene, Delay Of Germination 1 (DOG1), was found to influence not only germination, but also flowering time, overall life history, and fitness. Flowering time of the previous generation, in turn, imposed maternal effects that altered germination, the effects of DOG1 alleles, and the direction of natural selection on these alleles. Thus under natural conditions, germination genes act as flowering genes and potentially vice versa. These results illustrate how seasonal environmental variation can alter pleiotropic effects of developmental‐timing genes, such that effects of genes that regulate prior life stages ramify to influence subsequent life stages. In this case, one gene acting at the seed stage impacted the entire life cycle.     

Coexistence of two competing corixid species (Heteroptera) in an archipelago of temporary rock pools

Summary Coexistence of Callicorixa producta and Arctocorisa carinata in an archipelago of rock pools appears to be facilitated by division of the environment on the time axis. To test this, a hypothesis is built on r-K theory. As A. carinata is twice the size of C. producta it is expected that the former species is more K-selected living in more permanent pools where food resources are limiting and competition strong. The smaller C. producta is expected to prefer newly refilled pools with abundant food, and thus to be more r-selected. Gathered test data on 17 life history and population ecological features agree with the prediction. It is also shown that the both rock-pool corixids are more r-selected than other species of the family.     

Life history plasticity of Nemoura trispinosa (Plecoptera: Nemouridae) along a permanent-temporary water habitat gradient

1. The life history of the small herbivorous stonefly Nemoura trispinosa Claassen was studied in a variety of small springs in southern Ontario, Canada. Nymphs generally were able to tolerate a wide range of environmental conditions and were found in 78% of habitats sampled, although population densities differed markedly. 2. Life-cycle patterns varied from a univoltine, slow seasonal type to a univoltine, fast seasonal type with extended egg development. In one, highly stable, spring the life cycle was semivoltine. Inter-year variation was studied for 5 years in one spring and was found to be low relative to among-spring variation. 3. Differences in the life history traits of N. trispinosa populations from our spring series were most probably an expression of phenotypic plasticity rather than of genetic differentiation. 4. Maximum annual water temperature was the factor most influential on nymphal growth rate (non-linear relationship), whereas range in generation time was related to the degree of habitat permanence.     

The role of nutrition in regulation of predator-induced crests of Daphnia carinata

1. The effects of food level on the development of crests by Daphnia carinata in the presence of Anisops grains (Notonectidae) were investigated in two laboratory experiments and in the field. 2. In the laboratory experiments D. carinata were grown at different food levels either in the presence of A. gratus or in predator-free water. In the field study the effects of six variables (temperature, food level, density of Daphnia, density of Anisops, brood size and maternal nutrition) on development of crests in the next generation of Daphnia were investigated over a 12-month period at a single field site. 3. In the laboratory food level had a small but significant effect on relative crest height of D. carinata. In the presence of notonectid predators the rate of crest growth relative to body length was higher than for control D. carinata only during embryogenesis and/or the first two neonatal instars. Beyond this age the rate of crest growth was similar in both morphs. 4. In the field study, crest height was negatively correlated with density of Daphnia and positively correlated with density of Anisops and maternal nutrition (maternal nutrition was defined as the somatic weight of a female with a body length of 2.5 mm, calculated from the length-dry weight regression based on a sample of the population). However, these data were biased by a short period when no Anisops were present in the pond. When these data were excluded, the only factor which significantly correlated with crest height was maternal nutrition. This finding indicated that relative crest height of the next generation was determined primarily during embryogenesis on the basis of maternal feeding success and possibly on food availability during the first two instars.