Using morphological characters of subfossil daphniid postabdominal claws to improve taxonomic resolution within species complexes

Daphnia subfossils from lake sediments are useful for exploring the impacts of environmental stressors on aquatic ecosystems. Unfortunately, taxonomic resolution of Daphnia remains is coarse, as only a small portion of the animal is preserved, and so the identification of daphniid subfossils typically relies upon postabdominal claws. Daphniid claws can be assigned to one of two species complexes: D. longispina or D. pulex. Both complexes contain species with differing environmental optima, and therefore improved taxonomic resolution of subfossil daphniid claws would aid paleolimnological analyses. To identify morphological features that may be used to help differentiate between species within complexes, we used species presence/absence data from net tows to select lakes in central Ontario (Canada) containing only a single species from a particular complex, then used remains preserved in surface sediments of these lakes to isolate four Daphnia species: D. ambigua and D. mendotae from the D. longispina complex, and D. pulicaria and D. catawba from the D. pulex complex. Our analyses demonstrate that, within the D. longispina complex, postabdominal claw length (PCL) and spinule length can be used to distinguish D. mendotae from D. ambigua. In addition, within the D. pulex complex, there are differences between D. pulicaria and D. catawba in the relative lengths of the proximal and middle combs on the postabdominal claw. However, the number of stout spines on the middle comb is an unreliable character for differentiating species. Overall, our data demonstrate that greater resolution within Daphnia species complexes is possible using postabdominal claws; however, the process is arduous, and applicability will likely decrease with the number of taxa present.     

Daphnia diversity on the Tibetan Plateau measured by DNA taxonomy

Daphnia on the Tibetan Plateau has been little studied, and information on species diversity and biogeography is lacking. Here, we conducted a 4‐year survey using the barcoding fragment of the mitochondrial COI gene to determine the distribution and diversity of Daphnia species found across the Plateau. Our results show that species richness is higher than previously thought, with total described and provisional species number doubling from 5 to 10. Six of the taxonomic units recovered by DNA taxonomy agreed well with morphology, but DNA barcoding distinguished three clades each for the D. longispina (D. galeata, D. dentifera, and D. longispina) and D. pulex (D. pulex, D. cf. tenebrosa, and D. pulicaria) complexes. The sequence divergence between congeneric species varied within a large range, from 9.25% to 30.71%. The endemic D. tibetana was the most common and widespread species, occurring in 12 hyposaline to mesosaline lakes. The lineage of D. longispina is the first confirmed occurrence in west Tibet.     

Competitive dominance among Cladocera: Are single-factor explanations enough?

I examine several hypotheses for competitive dominance among zooplankton using data from 20 studies containing field and laboratory experiments on competition between cladoceran species. In threeDaphnia species from rockpools studied in the laboratory, the largeD. magna was the superior competitor at higher food levels and at 15°, while the smallerD. pulex andD. longispina were dominants at low food levels at 20°.D. pulex usually excludedD. longispina. No single-factor hypothesis gives a satisfying explanation for these results. A review of the literature data suggests that none of the following hypotheses for competitive dominance have sufficient support to be considered general: The size-efficiency hypothesis (supported in 60% of the tests), ther-max hypothesis (68% support), efficiency at low food levels (36% support), and that small species are superior competitors (only 17% support). Competitive ability and susceptibility to predation appear to be positively related (9 out of 10 cases). 76% of the experiments carried out under different environmental conditions showed varying outcomes. Competition between cladocerans should evidently be studied within a multi-factor framework. Since shifts in competitive advantage with environmental changes were common, the competitive exclusion principle may often not apply in zooplankton assemblages. Non-equilibrium coexistence of competitors in a variable environment, i.e. the paradox of the plankton, is a framework worthy of consideration in zooplankton, and possibly also in many other natural communities.     

Diel horizontal migration and swarm formation in Daphnia in response to Chaoborus

Dense swarms of Daphnia longispina (up to 4000 animals l^–1) were recorded along the littoral zone in a lake where Chaoborus flavicans is considered the main predator. D. longispina coexisted with D. pulex, but there were no D. pulex in the littoral swarms. Swarms were less dense at night (about 1/10 the density), and D. longispina exhibited diel horizontal migrations by aggregating in the littoral during the day and spreading out at night. Laboratory experiments showed that Chaoborus capture efficiency on juvenile daphnids was higher in the light compared to darkness, and that Daphnia exhibited a behavioural response to water that had previously contained Chaoborus. We conclude that predation from Chaoborus can be an important factor affecting the distribution patterns of Daphnia observed in this lake. The behavioural experiments indicated that this influence might be partly mediated by chemical agents.     

Variations of antioxidant enzymes in Daphnia species and populations as related to ambient UV exposure

To assess their role in photoprotection in the crustacean zooplankton Daphnia spp., activities of the antioxidants catalase (CAT), superoxide dismutase (SOD), glutathione transferase (GST) and content of carotenoids (CAR) were studied in 4 Daphniaspecies from a total of 50 populations. Included in the survey were alpine and lowland populations of both pigmented and non-pigmented D. longispina, rock-pool and laboratory clones of D. magna and Arctic populations of the D. pulex/D. middendorffianacomplex. Most of the surveyed populations inhabit shallow ponds, and are thus highly exposed to UV-radiation during the summer. The results are primarily discussed in relation to antioxidants as possible protection against UV-radiation. D. pulex, D. middendorffianaand D. longispinaoccurred both as hyaline and pigmented (carapace melanization) morphs. The most notable inter-species differences were the high activity of CAT in the D. magnalab-clone, the low activity of GST in melanic animals from the D. pulexgroup and the high activity of SOD in D. longispinafrom a lowland humic pond. Contrary to expectations, we found no differences in antioxidant capacity between melanic and non-pigmented alpine D. longispina. Intraspecies variation in antioxidant activities was studied in relation to pond characteristics. Among the alpine populations of D. longispina there was a significant positive correlation between absorbance (300 nm) of the pond water and CAT activity, which could be related to ambient levels of photoinduced hydrogen peroxide production in these small water bodies. The data from this comparative study provide insight in photo-protective mechanisms in cladocera, and constitute a valuable basis for future research on pro-oxidant and antioxidant processes in alpine and arctic freshwater zooplankton.     

CROSS‐SPECIES INFECTION TRIALS REVEAL CRYPTIC PARASITE VARIETIES AND A PUTATIVE POLYMORPHISM SHARED AMONG HOST SPECIES

A parasite's host range can have important consequences for ecological and evolutionary processes but can be difficult to infer. Successful infection depends on the outcome of multiple steps and only some steps of the infection process may be critical in determining a parasites host range. To test this hypothesis, we investigated the host range of the bacterium Pasteuria ramosa, a Daphnia parasite, and determined the parasites success in different stages of the infection process. Multiple genotypes of Daphnia pulex, Daphnia longispina and Daphnia magna were tested with four Pasteuria genotypes using infection trials and an assay that determines the ability of the parasite to attach to the hosts esophagus. We find that attachment is not specific to host species but is specific to host genotype. This may suggest that alleles on the locus controlling attachment are shared among different host species that diverged 100 million year. However, in our trials, Pasteuria was never able to reproduce in nonnative host species, suggesting that Pasteuria infecting different host species are different varieties, each with a narrow host range. Our approach highlights the explanatory power of dissecting the steps of the infection process and resolves potentially conflicting reports on parasite host ranges.     

Microsatellite and single‐nucleotide polymorphisms indicate recurrent transitions to asexuality in a microsporidian parasite

Assessing the mode of reproduction of microparasites remains a difficult task because direct evidence for sexual processes is often absent and the biological covariates of sex and asex are poorly known. Species with geographically divergent modes of reproduction offer the possibility to explore some of these covariates, for example, the influence of life‐history traits, mode of transmission and life‐cycle complexity. Here, we present a phylogeographical study of a microsporidian parasite, which allows us to relate population genetic structure and mode of reproduction to its geographically diverged life histories. We show that in microsporidians from the genus Hamiltosporidium, that use the cladoceran Daphnia as host, an epidemic population structure has evolved, most probably since the last Ice Age. We partially sequenced three housekeeping genes (alpha tubulin, beta tubulin and hsp70) and genotyped seven microsatellite loci in 51 Hamiltosporidium isolates sampled within Europe and the Middle East. We found two phylogenetically related asexual parasite lines, one each from Fennoscandia and Israel, which share the unique ability of being transmitted both vertically and horizontally from Daphnia to Daphnia. The sexual forms cannot transmit horizontally among Daphnia, but presumably have a complex life cycle with a second host species. In spite of the similarities between the two asexual lineages, a clustering analysis based on microsatellite polymorphisms shows that asexual Fennoscandian parasites do not share ancestry with any other Hamiltosporidium that we have sampled. Moreover, allele sequence divergence at the hsp70 locus is twice as large in Fennoscandian than in Israeli parasites. Our results indicate that asexual reproduction evolved twice independently, first in Fennoscandian and more recently in the Israeli parasites. We conclude that the independent origin of asexuality in these two populations is associated with the altered parasite mode of transmission and the underlying dynamics of host populations.     

Effect of Body Size on Toxicity of Zinc in Neonates of Four Differently Sized <Emphasis Type="Italic">Daphnia</Emphasis> Species

The sensitivity of neonates of four Daphnia species to zinc was tested in relation to their mean body size. These mean sizes of these four Daphnia spp were: D. magna, 0.813 ± 0.055 mm, D.␣pulicaria, 0.745 ± 0.063 mm, D. pulex, 0.645 ± 0.044 mm and D. galeata, 0.611 ± 0.058 mm. A positive relationship between EC₅₀ (24, 48) values and neonates size was found. The smaller the size of the daphnid the higher was the sensitivity to heavy metal toxicity. For all tested species did the EC₅₀ values decrease with time; the decrease was most marked for D. magna and the least for D. galeata. The EC₅₀ values of D. magna were higher than would be expected on basis of its body size.     

Microsporidiosis in a Daphnia pulex population

The impact of parasitism by Thelohania sp., an intracellular microsporidian, on Daphnia pulex inhabiting a vernal pond was studied for three years. Three issues were considered: the distribution of parasites among hosts, the effect of parasites on individuals, and the impact of parasitism on population growth. Each year, Thelohania infected members of at least half of the generations of Daphnia produced in the pond. When the frequency of infection was low, parasites were found only in large adults. As infection frequency increased, parasitism spread downward through host size classes. However, parasites were rarely found in juveniles. Parasitism reduced clutch size drastically, increased mortality to a variable extent, had little impact on egg size or on per instar growth and none on molt frequency. Interaction with other stresses, such as food limitation, exacerbated some of these effects. Parasitism reduced instantaneous birth rate much more than it elevated instantaneous death rate. Population growth was reduced significantly but it is unlikely that Thelohania alone regulates the growth of this Daphnia population.     

Two hundred years of a diverse Daphnia community in Lake Naivasha (Kenya): effects of natural and human-induced environmental changes

1. We used fossil diapausing eggs extracted from ²¹⁰Pb‐dated sediment cores to reconstruct historical changes in the Daphnia community of Lake Naivasha, a climate‐sensitive lake in Kenya which over the past 200 years has experienced a series of well‐documented natural and anthropogenic environmental changes. 2. Contiguous sampling and analysis of four cores yielded ephippial capsules of eight Daphnia species. Only two of these had been recorded previously in live collections from Lake Naivasha, and one species is new to science. The four more common species (Daphnia barbata, D. laevis, D. magna, and D. pulex) show striking differences in abundance patterns and population dynamics through time. Four other species (D. lumholtzi, D. curvirostris, D. longispina s.l., and Daphnia sp. nov. type Limuru.) appear to have been present only occasionally. Nevertheless, between 1895 and 1915 seven species of Daphnia inhabited Lake Naivasha simultaneously. 3. Despite considerable natural environmental change associated with climate‐driven lake‐level fluctuations, the Daphnia community of Lake Naivasha has been severely affected by human activities over the past century, especially the introduction of exotic fishes and water‐quality changes because of agricultural soil erosion. The recent reappearance of large‐bodied Daphnia species (D. magna, D. barbata, D. lumholtzi, Daphnia sp. nov. type Limuru) after 20–110 years of absence can be explained by their release from fish predation, following a dramatic increase in turbidity caused by excess clastic sediment input from eroded catchment soils. The small‐bodied species D. laevis has fared less well recently, presumably because the benefit of lowered predation pressure is counteracted by more pronounced negative effects of increased turbidity on this species and loss of submerged macrophyte beds which formerly served as predation refuge. 4. Our results suggest that, despite considerable environmental instability and the absence of specialised zooplanktivores, top‐down control of fish on large zooplankton is important in Lake Naivasha. Predation pressure from fish has led to clear‐cut shifts in local Daphnia species composition, but failed to drive the larger taxa to extinction.     

Spatial heterogeneity of daphniid parasitism within lakes

Spatially explicit models show that local interactions of hosts and parasites can strongly influence invasion and persistence of parasites and can create lasting spatial patchiness of parasite distributions. These predictions have been supported by experiments conducted in two-dimensional landscapes. Yet, three-dimensional systems, such as lakes, ponds, and oceans, have received comparatively little attention from epidemiologists. Freshwater zooplankton hosts often aggregate horizontally and vertically in lakes, potentially leading to local host–parasite interactions in one-, two-, or three-dimensions. To evaluate the potential spatial component of daphniid parasitism driven by these local interactions (patchiness), we surveyed vertical and horizontal heterogeneity of pelagic Daphnia infected with multiple microparasites in several north temperate lakes. These surveys uncovered little evidence for persistent vertical patchiness of parasitism, since the prevalence of two parasites showed little consistent trend with depth in four lakes (but more heterogeneity during day than at night). On a horizontal scale of tens of meters, we found little systematic evidence of strong aggregation and spatial patterning of daphniid hosts and parasites. Yet, we observed broad-scale, basin-wide patterns of parasite prevalence. These patterns suggest that nearshore offshore gradients, rather than local-scale interactions, could play a role in governing epidemiology of this open water host–parasite system. Electronic Supplementary Material Supplementary material is available for this article at     

Responses of algae, bacteria, Daphnia and natural parasite fauna of Daphnia to nutrient enrichment in mesocosms

Understanding responses of parasites to changes in nutrient regimes is necessary for prediction of their role in aquatic ecosystems under global change in nutrient loading. We studied the response of the natural parasite fauna of Daphnia longispina to nutrient enrichment in mesocosms in a small humic lake. We measured the concentrations of inorganic phosphorus and nitrogen in the water, total nutrients in the seston, algal and bacterial biomass, Daphnia population dynamics, Daphnia stoichiometry, Daphnia stable isotope values and the presence and abundance of parasites in treated mesocosms as compared to three control ones. Incorporation of the nutrient enrichment in the food web was seen as increased nutrient concentrations in the epilimnion and as a decrease in carbon:nutrient ratios and δ¹⁵N values in Daphnia. Nutrient enrichment did not significantly influence algal, bacterial or Daphnia biomass. One of the four parasite species observed, unidentified small gut parasite, had a higher prevalence (percentage of Daphnia infected) in treated mesocosms, but its intensity (number of parasites per infected host) remained the same among treatments. Our results suggest that the effect of nutrient enrichment on host–parasite dynamics is dependent on complex interactions within food webs and on the epidemiological traits of parasites.     

Population genetic structure of three pond-inhabiting Daphnia species on a regional scale (Flanders, Belgium)

1. Only a few studies have compared patterns of genetic variation among populations of different Daphnia species on a regional scale. The present study addresses this gap and examines the relationship between diversity as revealed by allozyme variation and habitat size for populations of Daphnia pulex, D. obtusa and D. curvirostris in Flanders (Belgium). In addition, we examined whether patterns of isolation‐by‐distance could be observed in each of these three Daphnia species. 2. The relationship between genetic diversity and habitat size varied among Daphnia species that occur in the same region. In D. pulex and D. obtusa populations, a positive relationship between local genetic diversity and habitat size was found, whereas the relationship was negative in D. curvirostris populations. 3. Regional genetic diversity was lower than expected from patterns of local genetic diversity in D. pulex and D. obtusa populations in Flanders. This suggests that the subdivision of local Daphnia populations in a region did not obviously increase genetic diversity. 4. Genetic differentiation among populations of these three species in Flanders was moderate and comparable with values observed in other Daphnia species. Patterns of isolation‐by‐distance could be observed, but the scatter was high (D. pulex) or the slope was very low (D. obtusa).     

Molecular genetic identification and phylogeny of Daphnia species (Crustacea, Cladocera) from water bodies of the Lake Chany basin

The data on the molecular genetic identification of Daphnia species from the water bodies of the Lake Chany basin are presented. Phylogenetic relationships between these species have been established. The fragments of the mitochondrial DNA genes were used as genetic markers. According to the data obtained, the water bodies examined were inhabited by five Daphnia species, including Daphnia (Daphnia) galeata Sars, D. (D.) longispina O. F. Müller, D. (D.) curvirostris Eylmann, D. (D.) pulex Leydig, and D. (Ctenodaphnia) magna Straus. In addition, longispina a-like individuals that form a separate phylogenetic lineage was identified.     

Size and shape ofDaphnia longispina in rock-pools

The body size and shape ofD. longispina in small and large rock-pools was measured. The mean body length ofDaphnia in large rock-pools with vertebrate planktivores was smaller than that ofDaphnia in small rock-pools without vertebrates, but the variability in body lengths within pools over the season was as great as that found between pools and predator regimes. We did find thatD. longispina in large rock-pools produced one egg at a smaller body length and had fewer eggs per individual than didDaphnia in small rock-pools.D. longispina populations also showed different body shapes both as the season progressed and in rock-pools having different predator composition. As expected inDaphnia cyclomorphosis, the core body length decreased during the summer, accompanied by significantly more pointed heads and longer tail spines than in early summer.D. longispina in large rock-pools (with vertebrate predators) have significantly larger exuberances than those in small rock-pools (lacking vertebrate planktivores). The results are discussed in the context of size-selective predation.     

Daphnia community analysis in shallow Kenyan lakes and ponds using dormant eggs in surface sediments

1. Water fleas of the genus Daphnia are considered rare in tropical regions, and information on species distribution and community ecology is scarce and anecdotal. This study presents the results of a survey of Daphnia species distribution and community composition in 40 standing waterbodies in southern Kenya. The study sites cover a wide range of tropical standing aquatic habitats, from small ephemeral pools to large permanent lakes between approximately 700 and 2800 m a.s.l. Our analysis combines data on Daphnia distribution and abundance from zooplankton samples and dormant eggs in surface sediments. 2. Nearly 70% (27 of 40) of the sampled waterbodies were inhabited by Daphnia. Although their abundance in the active community was often very low, this high incidence shows that Daphnia can be equally widespread in tropical regions as in temperate regions. 3. Analysis of local species assemblages from dormant eggs in surface sediments was more productive than snapshot sampling of zooplankton communities. Surface‐sediment samples yielded eight Daphnia species in total, and allowed the detection of Daphnia in 25 waterbodies; zooplankton samples revealed the presence of only four Daphnia species in 16 waterbodies. 4. Daphnia barbata, D. laevis, and D. pulex were the most frequently recorded and most abundant Daphnia species. Canonical correspondence analysis of species–environment relationships indicates that variation in the Daphnia community composition of Kenyan waters was best explained by fish presence, temperature, macrophyte cover and altitude. Daphnia barbata and D. pulex tended to co‐occur with each other and with fish. Both species tended to occur in relatively large (>10 ha) and deeper (>2 m) alkaline waters (pH 8.5). Daphnia laevis mainly occurred in cool and clear, macrophyte‐dominated lakes at high altitudes.     

Intraspecific trait variation and colonization sequence alter community assembly and disease epidemics

When individuals from multiple populations colonize a new habitat patch, intraspecific trait variation can make the arrival order of colonists an important factor for subsequent population and community dynamics. In particular, intraspecific priority effects (IPEs) allow early arrivers to limit the growth or establishment of later arrivers, even when competitively inferior on a per‐capita basis. Through their effects on genes and traits, IPEs can alter short‐term growth and long‐term evolutionary change in single species metapopulations. Given their importance for intraspecific interactions, IPEs in a dominant species have the potential to affect the composition of entire communities. We conducted an experiment to determine whether and how arrival order and IPEs in the zooplankter Daphnia pulex affected its interactions with both competitors (the cladoceran Simocephalus vetulus) and parasites (the virulent fungus Metschnikowia bicuspidata). We found strong evidence for IPEs in Daphnia, as early arrivers inhibited late arrivers even when competitively inferior. These IPEs in Daphnia altered both the establishment success of interspecific competitors and the size of disease epidemics: early colonization by fast‐growing D. pulex led to large Daphnia populations and low competitor establishment, but large disease epidemics. Early colonization by slow‐growing D. pulex, on the other hand, resulted in small Daphnia populations with high competitor establishment, but smaller disease epidemics. Overall, our results demonstrate the importance of intraspecific variation and arrival order for community dynamics, and highlight IPEs as a general mechanism driving variation in natural communities.     

Morphological defenses induced in situ by the invertebrate predator Chaoborus : comparison of responses between Daphnia pulex and D. rosea

The presence of plankton predators may induce altered morphology in their potential prey. To date, the mechanism of induction and adaptive value of such defensive responses have been examined in the laboratory. This study investigated the morphological defense structures induced by the invertebrate predator Chaoborus in two coexisting Daphnia species, D. pulex and D. rosea, in the field. In Piscivore Lake (Gr?fenhain, Germany), continuous and intense biomanipulation had led to near elimination of planktivorous fish and greatly increased abundances of Chaoborus (up to >10 larvae l^–1). Here, the density of Chaoborus was manipulated within the lake by an enclosure/exclosure setup and resulting morphological responses of Daphnia spp. were investigated in situ. Three replicate enclosures (4.6 m³) contained no Chaoborus (predator exclusion bags), whereas Chaoborus entered three others at ambient densities (predator enclosures). In both species of Daphnia, formation of neckteeth and elongation of the tail spine were recorded in the predator enclosures, but not in the predator exclusion treatments. Additionally, D. rosea responded to predator inclusion with an increase of the size at first reproduction. Despite the induced defense structures, the presence of Chaoborus caused increased mortality of both Daphnia species. In addition, Chaoborus affected the coexistence of the two populations of Daphnia by causing higher relative mortality in D. rosea. Neckteeth formation was always more pronounced in D. pulex than in D. rosea of the same size. Neckteeth were induced specifically in vulnerably sized juvenile instars of D. pulex, but were not found in all vulnerable instars of D. rosea. In D. rosea, neckteeth were few or absent in the ephippial hatchlings, and neckteeth formation ceased before juveniles reached a body size outside the range that larger larval stages of Chaoborus could ingest. This study provides the first experimental demonstration in the field of the inducibility of morphological defense structures in Daphnia at ambient densities of Chaoborus larvae, and quantifies these in situ responses. This expands on earlier observations of a correlation between predator density in the field and the expression of neckteeth in Daphnia. The term ”maximum size for neckteeth formation” (MSNF) is defined as the limit in body size above which no production of neckteeth was evident. This limit was used to distinguish the size classes of Daphnia that show a sensitive response to Chaoborus kairomone. This new term may be used for further comparisons among species and among different types of predator-induced responses as well as for the evaluation of the adaptive value of defense structures. Received: 10 April 1999 / Accepted: 6 April 2000     

Changes in epibiotic burden during the intermolt of Daphnia determined by hypodermal retraction stages

We calibrated four stages of hypodermal retraction and cuticle regeneration with five stages of parthenogenetic egg development in Daphnia. Using the hypodermal retraction stages, we found that epibiotic burden increased with elapsed intermolt time for juvenile, male, and female Daphnia bearing parthenogenetic or ephippial eggs. The rate of increase of burden was similar for adult females of two Daphnia species and for males and females of D. pulex. Rate of increase of burden was similar between juvenile and adult females of D. galeata mendotae and D. pulex.     

Somatic polyploidy examined by flow cytometry in Daphnia

Flow cytometry was used to examine the patterns of somatic polyploidyand to determine the haploid genome sizes in Daphnia. The averageproportions of polyploid nuclei among adult animals of D.pulex,D.longispina, D.pulex x D.longispina and D.magna equalled 27,24, 24 and 24%, respectively. Both interclonal and developmentallyregulated variation were observed in the level of somatic polyploidy.Adult animals expressed more extensive polyploidy than did juveniles.The estimates of the haploid genome sizes (C values) of D.pulex,D.longispina, D.pulex x D.longispina and D.magna equalled 0.32,0.27, 0.26 and 0.37 pg, respectively. These are the smallestgenomes reported in crustaceans. Apparently, somatic polyploidycompensates for the loss of genetic material and gives riseto genetic plasticity which may contribute to the considerablelevel of phenotypic plasticity observed in Daphina.