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.     

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 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.     

Sexual reproduction in Daphnia: interspecific differences in a hybrid species complex

To investigate whether reproductive variation in the Daphnia galeata-cucullata-hyalina hybrid species complex can explain the occurrence of hybrids and backcrosses, a total of 43 clones from three north German lakes were tested for allocation to sexual reproduction under equal stress conditions. Six replicates per clone were followed until the seventh adult instar. I used the following cues to promote sexual reproduction: short photoperiod, water from a crowded Daphnia culture, fish-conditioned water and low food concentration. For each animal, clutch size and clutch sex were recorded. Ephippia, which were empty since the animals were cultured individually, were only produced by D. cucullata and D. cucullata x hyalina (26% and 6% of the broods, respectively), whereas almost all taxa produced males (the range was 2–15%). Intraspecific variation for male and ephippial production was also found. The fact that the taxa tested show different responses to the stimuli of-fered may indicate that there is reproductive isolation between them. This is supported by field data from the Tjeukemeer (The Netherlands) which shows that D. galeata mostly produces ephippia in spring and D. cucullata in autumn. The presence of hybrids with ephippia in both seasons, however, shows that backcrossing is also possible.     

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.     

Morphological differentiation within the Daphnia longispina group

Although morphological evolution is assumed to be slow within Daphnia species complexes, discontinuities in morphological space can be detected. Here, morphological data derived from females of genetically-defined clones (cf. Gießler et al., 1999) are presented, in order to estimate the genetic component of phenotypic variance under standardised laboratory conditions. Animals originated from clonal assemblages of pre-alpine lakes and ponds, and a remote lake in western Germany, covering a wide range of morphotypes known from the traditional species D. cucullata, D. galeata, D. hyalina, D. rosea, and a variety of interspecific hybrids. Phenotypic analyses were based on quantitative and qualitative morphological characters of females in the first and fifth instars. Morphological divergence between clones was analysed using discriminant analysis or multidimensional scaling and the significance of the morphological groupings was estimated using neighbour-joining trees and bootstrapping. All analyses confirmed that (a) phenotypic similarities among taxa change with instar, (b) in contrast to low genetic divergence, pronounced morphological divergence exists between animals separated on the lake/pond level favouring speciation by the habitat shift hypothesis.     

An investigation of allelopathic effects of Daphnia

• 1 The hypothesis is tested that large daphnids are able to suppress their own and other species' feeding and reproduction by means of excreting an inhibitory chemical (or chemicals).
• 2 In laboratory experiments with an Australian species, Daphnia carinata, water preconditioned with 3–67 daphnids 1-^?1 for 30 h had the effect of reducing feeding rates of D. carinata and D. lumholtzi provided with Selenastrum capricornutum.
• 3 For the two Daphnia species, there were highly significant negative correlations between feeding rate and the preconditioning density of D. carinata.
• 4 Water preconditioned with 20–30 daphnids 1-^?1 for 1–2 weeks reduced the grazing rates of Daphnia, Moina, and Diaphanosoma 2–3-fold.
• 5 Moina kept in such water for 2 days stopped feeding. Conditioned water kept for 3 days without animals still inhibited grazing by Moina. Hearing to 100°C removed the inhibitory effect.
• 6 Given excess food, and in non-renewed water, a gradient of D. carinata densities developed a strong negative correlation between clutch size and daphnid density after a 6-day time lag. This result may help explain the direct density-dependent regulation of cladoceran reproduction observed earlier in a subtropical lake.

     

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.     

Analysis and comparison of a set of expressed sequence tags of the parthenogenetic water flea <Emphasis Type="Italic">Daphnia carinata</Emphasis>

The water flea Daphnia carinata (D. carinata) reproduces both sexually and parthenogenetically, yet little is known about the genes involved in these processes. To further clarify the reproductive biology of Daphnia and elucidate their unique mechanism of reproductive transformation, we have generated and characterized an expressed sequence tag (EST) data set from D. carinata. A set of 1,495 clusters were generated from sequencing 3,072 randomly chosen clones from a parthenogenetic, juvenile water flea cDNA library. The nucleic acid and deduced amino acid sequences were compared with known GenBank sequences. Functional annotation found that 959 clusters showed significant homology with known genes involved in a broad range of activities, including metabolism, translation, development and reproduction, as well as genes involved in sensing environmental factors. We speculate that genes involved in development and reproduction, along with genes that allow the organism to sense changes in the environment, play important roles in the process of parthenogenetic reproduction and could be markers of the early steps of sexual differentiation. Additionally, 86% of the D. Carinata unique sequences could be stringently mapped to the D. pulex genome, of which 125 mapped to intergenic and intronic regions on the current assembly. Our results provide practical insight into crustacean reproductive biology, in addition to establishing a new animal model for reproductive and developmental biology. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users. Xiaoqian Xu and Shuhui Song contributed equally to this work. Nucleotide sequence data reported are available in the GenBank databases under the accession numbers: GD269049−GD272045.     

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.     

Field and laboratory studies on Daphnia carinata King (Cladocera: Daphnidae) from a seasonal tropical pond

The life history and seasonal variation in egg production of Daphnia carinata King, with reference to the seasonal variation in physico-chemical as well as biological factors of a tropical temporary pond, were studied. Laboratory observations on the life history of D. carinata were also made. The seasonal events of D. carinata are discussed in relation to recent studies on Cladocerans.     

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.     

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.     

Intra-specific rDNA-ITS restriction site variation and an improved protocol to distinguish species and hybrids in the Daphnia longispina complex

A collaborative research effort was undertaken to evaluate the robustness of a recently developed genetic tool for species identification of members in the morphologically variable Daphnia longispina species complex. This genetic method, based on restriction fragment length polymorphism (RFLP) of the internal transcribed spacer region (ITS) of nuclear ribosomal DNA (rDNA) with restriction enzymes Mwo I and Sau96 I [Billiones et al., 2004. Hydrobiologia 526: 43–53], was applied to many different European populations. Results were compared with two or more independently obtained characters (morphology, allozymes, mitochondrial DNA (mtDNA), or cloned rDNA-ITS sequences). Individuals of most taxa were readily identified, but unexpected ITS-RFLP patterns were found in many individuals indicated by other markers to be D. galeata or one of its hybrids. Among 43 investigated D. galeata populations (902 specimen analysed by ITS-RFLP), deviant RFLP fragment patterns occurred in 26 (i.e., more than half) of the populations. The deviant patterns could be attributed to the loss of one single restriction site in the ITS2 region. This loss made the distinction of D. galeata from other species unreliable, and F1 hybrids could not be identified. Future users should be aware of this shortcoming of the Billions et al. [2004. Hydrobiologia 526: 43–53] protocol. As a solution to this problem, we present an improved genetic identification protocol based on a simple double digestion of the rDNA-ITS region with the restriction enzymes BsrB I and EagI. Sequence analyses of rDNA-ITS clones and preliminary testing indicate that the new protocol is unaffected by the rDNA variation which troubled the Mwo I/Sau96 I protocol. Further, the new protocol identifies all European species of the D. longispina complex, as well as their F1 hybrids. However, a wider screening is required to verify its general utility for all species, since yet unknown variation may occur. Guest editor: Piet Spaak Cladocera: Proceedings of the 7th International Symposium on Cladocera     

Systematic analysis of Dendrobium Swartz section Dendrocoryne in the Australian region

Dendrobium sect. Dendrocoryne is a difficult taxonomic group of allied species and species complexes, occurring on the eastern Australian coastline, Lord Howe Island and New Caledonia. Significant morphological characters were assessed from all key areas of the distribution range. Patterns of variation between species, varieties and hybrids were investigated by principal coordinate analysis. The analyses affirm 12 species in the sect. Dendrocoryne described by Schlechter (1912, 1982) and Dockrill (1969, 1992), including a cluster of small statured D. gracilicaule, D. adae, D. fleckeri, and D. finniganense, and three variable species complexes – D. speciosum, D. kingianum and D. tetragonum. Recently described D. finniganense and D. callitrophilum are interpreted as members of the section. Phylogenetic relationships are presented based on parsimony analysis of 31 morphological characters. Analyses do not support proposals to recognize new genera for D. tetragonum, D. callitrophilum and D. aemulum, or to create new species within D. tetragonum, D. jonesii, D. speciosum and D. kingianum. The phylogeny indicates that rainforest taxa are earlier lineages and xerophytic taxa are more derived.     

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     

Tumbleweed (Salsola, section Kali) species and speciation in California

Tumbleweeds (Salsola species, section Kali) are road side and rangeland pest plants throughout the 48 contiguous states in the US. Three described tumbleweed species and two undescribed Salsola taxa occur in California. The known species are Russian thistle, Salsola tragus, introduced from Eurasia in the 1800s, Russian barbwire thistle, S. paulsenii, which grows in the desert regions of California, and is also native to Eurasia, and the recently identified S. kali subspecies austroafricana, possibly native to South Africa. Our goals were to investigate karyology, genome size, and molecular genetic affinities of the described species and the other taxa within their ranges in California using recently developed microsatellite loci, dominant nuclear DNA markers (RAPD and ISSR), and DNA sequence data. Chromosome counts and genome size assessments made with flow cytometry were compared. These analyses indicated that one undescribed taxon is a new allopolyploid hybrid between S. tragus and S. kali subspecies austroafricana, and the other undescribed taxon appears to be a complex hybrid involving all three described species. The invasion potentials for the hybrid taxa are unknown. Tumbleweeds are the focus of biological controls efforts but the identification of suitable agents for the hybrid taxa may be problematic because of the large amount of genetic variability encompassed within this evolving Salsola complex.     

Direct observation of toxic effects of cyanobacterial extracellular products on Daphnia

The filtrate from a suspension of the cyanobacterium Anabaena minutissima var. attenuata depressed thoracic limb beat frequency of Daphnia carinata by 38–45% in a food-free medium. Repeated exposure to the filtrate produced a similar depression of activity with full or neary full recovery. Response time to the filtrate was 5–8 min and recovery time was 8–12 min. The dose effect on limb beat frequency was continuous, linear, correlated with increased concentration and with no threshold. There was no relationship between body length and limb beat frequency.The interaction between toxicity and food concentration was tested using the diatom Cyclotella suspended in Anabaena filtrate. Daphnia limb beat frequency was depressed by 62%.