Cyclomorphosis in Daphnia cucullata: morphometric and population genetic analyses

The cyclomorphic change of head and tailspine of Daphnia cucullatahas been analysed morphometrically and compared with temporalanalyses of enzyme variability. During June, round-headed animalsare rapidly replaced by specimens with high helmets; the rateof increase of the relative helmet length is more rapid thanthe rate of decrease later in the year. The relative sizes ofhelmet and tail spine are not always coupled. The tailspinelength remains approximately the same year round and grows withnegative allometry, so that small animals have the relativelylongest spines. In contrast, the helmet grows in a positivelyallometric fashion and exhibits high seasonal variation. Helmetgrowth is strongly correlated with water temperature but notwith food abundance. The morphometric results are consistentwith the hypothesis that helmet formation is a response to fishpredation while the tail spine provides protection against invertebratepredators. There is no indication of changes in the gene orgenotype frequencies at two loci during the period of rapidmorphological change. Thus cyclomorphosis apparently appearsin the absence of clonal replacement.     

Cyclomorphosis as a factor explaining success of a Daphnia hybrid in Tjeukemeer

Recent studies have shown that Daphnia hybrids are intermediate to their parental species for several life history traits, and that they combine advantageous traits of the parental species, thereby decreasing their risk of being preyed upon by planktivorous fish. In this study D. galeata, D. cucullata and their interspecific hybrid are compared with respect to cyclomorphosis and juvenile growth as a potential mechanism to avoid invertebrate predation. Helmets and tail spines are known to reduce predation risk, whereas fast juvenile growth reduces the period that Daphnia are vulnerable to predation. In laboratory cultures, I tested the hypothesis that, with regard to cyclomorphosis, juvenile hybrids are more comparable to the juveniles of D. cucullata, and with respect to growth rate, they are more comparable to D. galeata. The hybrid did not differ significantly from D. galeata in the time needed to reach 0.9 mm body length. In contrast, the relative spine and helmet length of juvenile hybrids were more comparable with those of D. cucullata, whereas in adult instars these characters were more similar to D. galeata. Thus during the first juvenile instars, the hybrids combine the relative long helmet and spine of D. cucullata, with the faster growth of D. galeata. This combination of characteristics might reduce hybrid vulnerability to invertebrate predation in field situations.     

Induction of helmet development by a Chaoborus factor in Daphnia ambigua during juvenile stages

Daphnia ambigua juveniles developed spike-like helmets inducedby a water-soluble factor from Chaoborus, when reared in Chaoborusflavicans-conditioned medium. A study was made of the time atwhich the Chaoborus factor affects helmet development. Juvenilesdeveloped the helmets when individuals were exposed to the Chaoborusfactor after being born, but did not develop the helmets whenthe individuals were exposed to the factor only in the egg orembryo stages. These results indicate that D.ambigua respondsto the Chaoborus factor during the juvenile stages.     

Daphnia invasion: population dynamics of Daphnia assemblages in two eutrophic lakes with particular reference to the introduced alien Daphnia ambigua

The population dynamics of Daphnia species (D.cucullata, D.ambigua,D.galeata) in two small, productive lakes were studied and experimentswith field enclosures were carried out to elucidate the ecologicalniches of the introduced, alien species D.ambigua. Daphnia ambiguocoexisted temporally with the similarly sized D.cucullata, butlived in deeper water. Although both species had the same clutchsizes, similar proportions of ovigerous females, and death rates.D.cucullata was nurnerically dominant over D.ambigua. This suggeststhat the observed difference in abundance is simply a resultof the slower population growth rates of D.ambigua at lowertemperatures in greater depths. The experiments with field enclosuresshowed that when D.culcullata and D.ambigua were forced to livein the same layer of water, D.ambigua was numerically dominantover D.culullata both in the enclosures with and without fish.As the difference between the abundances of the two speciesdecreased in the enclosure with fish, D.cucullata seems to bebetter protected against fish predation than D.ambigua. In theenclosures without fish, D.ambigua and D.cucullata were numericallysurpassed by the larger species D.galeara. The present resultsimply that in some productive lakes with a high fish stock,D.mbigua invaded the niche of D.galeara and that the speciescan become an important component of the native zooplankton.The results also support the opinion that co-existing Daphniacan have very similar dynamics and niches. ¹ Present address: Zoologisches Institut, Christian-Albrechts-UniversitätKiel, Olshausenstrasse 40, D-24098 Kiel, FRG     

Cyclomorphosis in Daphnia lumholtzi induced by temperature

• 1 Cyclomorphosis is a well known phenomenon in Daphnia that involves a regular, seasonal, or induced change in body allometry. Long helmets and tail spines were induced in laboratory cultures of Daphnia lumholtzi with temperature of 31 °C as the proximal cue (temperature of locally occurring peak abundance in Kentucky Lake). The effect was greater in embryos than juveniles or adults exposed to the temperature cue.
• 2 The temperature cue appears to have a threshold value (animals cultured at 25 or 28 °C did not develop elongated helmets or spines). The helmet and spine length receded both with D. lumholtzi kept at a constant 31 °C temperature and when water temperature was decreased.
• 3 The induced helmet in this experiment (0.66 mm, 1.0 mm animal) was significantly longer than values reported in the literature for induction by planktivorous fish kairomones (0.25 mm, 1.2 mm animal). The strong response to a proximal cue of temperature may require the second weaker chemical cue for maintenance. It is suggested that a synergistic explanation with two cues may be more appropriate for cyclomorphosis induction and maintenance in Daphnia lumholtzi that could be tested with further studies.

     

Instar-dependent mortality rates of coexisting Daphnia species in Lake Vechten, The Netherlands

In Lake Vechten, population parameters were determined for twocoexisting Daphnia species, D.hyalina and D.cucullata, fromApril 23 to December 10, 1986. Daphnia hyalina is dominant inspring and D.cucullata in summer and autumn. Size frequencydistributions were converted into instar frequency distributionsby using experimental growth curves and field measurements onnewborn size and the size at first reproduction. A discreteevent computer model was used to calculate the instar mortalityrates. Relative importances of instar mortality rates for thepopulation mortality rate were calculated and discussed. Themortality of the juvenile stages was found to be more importantthan adult mortality during most of the sampling season, despiteegg mortality when egg-bearing females are killed. Only in Septemberand October was the mortality of adult instars more important.Correlation coefficients between differences in the rates ofincrease and differences in mean brood size and instar mortalitiesshow that the former are caused mainly by differences in juvenilemortality, the main factor determining the replacement of D.hyalinaby D.cucullata during early summer.     

Cyclomorphosis in Daphnia galeata mendotae Birge and D. retrocurva Forbes as a predator-induced response

SUMMARY. 1. Populations of Daphnia galeata mendotae and D. retrocurva were exposed for one generation to predators enclosed in mesh bags. The exposed Daphnia showed adaptive morphological changes that mimic cyclomorphosis in nature. The invertebrate predators Chaoborus and Notonecta induced longer helmets, while the fish Lepomis induced shorter bodies but had no effect on helmet length. Longer tail spines (relative to body length) were induced by Notonecta and Lepomis.
2. The responses of D. retrocurva were influenced by algae concentration, with the more extreme responses occurring at a higher food concentration and higher lipid index.     

The adaptive significance of cyclomorphosis in Daphnia: more possibilities

SUMMARY. Morphological variability in Daphnia populations has often been uncritically ascribed to phenotypic plasticity. For instance, detailed study revealed that the 'cyclomorphic species' D. carinata s. l. was a complex of nine species. Several of these species often cohabit and seasonal change in their relative frequencies causes phenotypic cycles which mimic true cyclomorphosis. Intraspecific genetic variation in head shape also seems widespread and is likely to be important in explaining phenotypic changes in many single species populations.
The hypothesis that helmet formation in Daphnia is primarily related to predator avoidance is not supported by work on the D. carinata group. Seasonal trends in species composition can be explained without reference to differential predation. Natality differences exist between species with disparate head size suggesting that helmet formation may have direct effects on fitness. Two possibilities are considered. The length of the anterior adductor muscle is directly correlated with helmet size and such variation may affect swimming efficiency. In addition the laminar design of helmets suggests a role in gas exchange.     

Instar-specific mortalities of coexisting Daphnia species in relation to food and invertebrate predation

Instar-specific mortalities of Daphnia hyalina and D.cucullatawere studied from May 19 to September 29, 1988 in combinationwith invertebrate predator and phytoplankton abundance. Simultaneouslife-table experiments were conducted under semi-natural conditionsin the laboratory to estimate juvenile mortality in a predator-freeenvironment. Juvenile mortality by predation was calculatedas the difference between juvenile mortality in the field andin the experiments and was the most important factor for thedifferences in abundance of the two species. For D.hyalina juvenilemortality was higher in early summer and probably caused byselective predation by Chaoborus flavicans. Predation by Leptodorakindtii was probably more important during the rest of the summer.Estimated mortality by predation adequately explained juvenilemortality, except for a 3-week period in August. Decreasingflagellate densities in July were accompanied by increased juvenilemortalities of D.hyalina and D.cucullata in the life-table experimentsin August and coincided with a Daphnia population decrease.     

The cyclomorphic response of Daphnia galeata mendotae: polymorphism or phenotypic plasticity

This study tests the hypothesis that cyclomorphosis in Daphniagaleata mendotae is equated with seasonal changes in the geneticstructure of populations. Multilocus electrophoretic genotypingof individual D. g. mendotae was used to identify related parthenogeneticlines. Between different clones and genotypes, there was phenotypicvariability in both laboratory and field populations. Withinclones and within genotypes, and in the absence of intense invertebratepredation, phenotypic expression was conservative. These patternssuggested that there was a correlation between phenotypic andgenetic change. However, in the field, only two of three populationsshowed seasonal changes in genotypic frequency, whereas allthree showed seasonal changes in phenorype. Furthermore, thepatterns of phenotypic change seen in the field populationswere different and appeared to be correlated with changes inphysical conditions and predation pressure. We conclude thathelmet morphology within genotypes is plastic enough to accountfor much of the seasonal variability observed in three populationsof D. g. mendotae and that genotypic frequency changes are notnecessary to explain seasonal changes in D. g. mendotae phenotype.     

Spatial and temporal patterns of sexual reproduction in a hybrid Daphnia species complex

Evidence for extensive interspecific hybridization among speciesof the genus Daphnia has been accumulating on a global scale.Although there is evidence for limited gene flow between taxavia hybridization, many species still maintain discrete morphologicaland molecular characteristics. We studied temporal and spatialpatterns of sexual reproduction within the Daphnia galeata–hyalina–cucullatahybrid species complex in a lake (Plußsee), located innorthern Germany. Allozyme electrophoresis allowed us to trackseasonal changes in taxon composition as well as the quantificationof back-crosses. Sexually-reproducing animals (ephippial femalesand males) were mainly found in autumn. The simultaneous presenceof sexual morphs of D. galeata and D. galeata x hyalina withthe dominant D. hyalina taxa makes recent hybridization, aswell as back-crossing, plausible. Males and ephippial femalesof D. hyalina were not back-crossed as were the parthenogeneticfemales. The low number of sexual clones of the hybrid D. galeatax hyalina might reflect its reduced fertility, although thesefew clones were detected in high densities. Only hybrid-clonesthat had a back-cross genotype (towards D. hyalina) exhibitedephippial females and males. This indicates that male and ephippialfemale production within the Daphnia taxa is not random, whichmight increase the chance for the parental Daphnia species toremain distinct.     

Morphological and mitochondrial-DNA analysis of the Indo-West Pacific rock oysters (Ostreidae: Saccostrea species)

Species of Saccostrea are common oysters on Indo-Pacific rockyshores. The taxonomy of this genus is unclear, however, becauseof morphological plasticity among its constituent species. Oystersamples with distinct morphotypes were collected from shoresexperiencing different degrees of wave exposure in Japan, Taiwan,China, Vietnam, Singapore and Australia. Intraspecific and interspecificvariations in shell characters have been identified. Only S.kegaki could be readily differentiated from S. cucullata-B,S. cucullata-C and S. mordax by multivariate shell allometricanalyses. Phylogenetic analyses using partial mitochondrial16S DNA sequences of approximately 513 base pairs identifiedtwo clades that correspond to superspecies, i.e. putative S.cucullata and putative S. mordax, comprising nine and two lineagesfrom the Indo-West Pacific, respectively. These are S. cucullata-A,B, C, D, E, F, G, S. kegaki and S. glomerata in one clade, andS. mordax-A and B in the other. Interspecific sequence divergence(Kimura-2-parameter values) between lineages ranged from 1.02%to 11.74%. Both S. cucullata-C and S. kegaki are restrictedto the northern hemisphere, whereas S. glomerata seems to bea temperate species from the southern hemisphere. S. cucullata-Doccurs only in Taiwan. Other species, i.e. S. cucullata-A, B,D, E and F and S. mordax-A and B, have a wide range on rockyshores in the tropical and subtropical Western Pacific and inboth northern and southern hemispheres. (Received 7 January 2005; accepted 4 November 2005)     

Environmental cues for cyclomorphosis in Daphnia retrocurva Forbes

SUMMARY.

• 1 The invertebrate predators. Chaohorus, Leptodora, Hydra, Utricularia, and cyclopoid copepods were ineffective as inducers of high helmets in laboratory reared Daphnia retrocurva. Lako water from different sources, when used as culture medium, was also ineffective.
• 2 High helmets were induced when clones were grown under warm and turbulent conditions. Presence or absence of illumination and varying photoperiod did not affect helmet growth in warm turbulent cultures. Different clones grown under warm turbulent conditions differed in their responsiveness to the treatment, but no relationship between helmet size in clones and helmet size in the field on the dates of clone isolation was demonstrated. Laboratory-induced phenotypes were rarely as extreme as those of field animals.

     

Morphological responses of Daphnia ambigua to different concentrations of a chemical extract from Chaoborus flavicans

1. Daphnia ambigua were reared individually with eight different concentrations of a chemical extract from larvae of Chaohorus flavicans, and the frequency of appearance of helmeted individuals and the helmet sizes were determined in the instars 1–6. 2. The animals developed most marked helmets in the second instar in response to the extract. The frequency of the helmeted individuals and helmet size in the second instar increased with increasing chemical concentration, although their dose-response curves differed between them. 3. The results may explain the cyclomorphosis of Daphnia, which develop the highest helmets in summer, when the predators are abundant and active, and the concentrations of the predator-released chemicals would presumably be high. 4. The extract became toxic to Daphnia at the highest concentration prepared.     

Resource depression in Daphnia galeata, Daphnia cucullata and their interspecific hybrid: life history consequences

In this study, the effect of the food concentration on severallife history parameters of Daphnia galeata, Daphnia cucullataand the hybrid between these two species was examined. The foodconcentration had a significant effect on both growth and reproductionof all three taxa. Mortality was highly species specific, andto a lesser extent dependent on the food concentration. Changesin life history parameters resulted in changes in the intrinsicrate of population increase (r). The food dependence of thedifferent species was found to be different. Daphnia cucullatahad a relatively constant and low r. The r values for the hybridvaried most, with low values compared with the parental specieswhen grown at low food levels and relatively high values athigher food concentrations. Our results suggest that when foodconcentrations are high the hybrid will have a selective advantageover both parental species, but that at lower food levels D.galeatawill have an advantage over both D.cucullata and the hybrid.     

Daphnia populations in three interconnected lakes with roach as the principal planktivore

Data on two co-existing Daphnia species, D.cucullata (G.O.Sars)and D.hyalina (Leydig), from three neighboring and interconnectedlakes in northern Poland, revealed an extremely confined rangeof population density fluctuations throughout summer in eachof the two species in each of the three lakes, with D.cucullatabeing an order of magnitude more abundant than D.hyalina (meanof 11.5 ind. l^–1 for D.cucullata and 1.1 ind. l^–1for D.hyalina with 95% confidence limits of ±2.5 and±0.5 ind. l^–1, respectively), in spite of markedlychanging fecundity. There was no apparent phase of decline orincrease that could be related to the distinct phases of lowor high fecundity resulting from different food levels. Analysisof the gut contents of roach (Rutilus rutilus), the dominantplanktivore in the three lakes, revealed identical selectivityfor each of the two Daphnia species, suggesting that the lowerdensity of the D.hyalina population was compensated for by thegreater conspicuousness of individuals of this species. It isconcluded that the population density of these Daphnia speciesremains far below the carrying capacity of the habitat and doesnot depend on food levels. Food availability merely sets therate of population increase, while the actual population densityin the lakes studied reflects the species' vulnerability topredation by planktivorous fish.     

Cyclomorphosis in Daphnia: an adaptation to avoid invertebrate predation

Seasonal morphological changes in three Daphnia species were followed over a two-year period in two lakes that differ in invertebrate and fish pressure. Whereas the morphology of D. hyalina, the biggest of the three species, varied little from season to season, D. cucullata, the smallest, exhibited the most pronounced seasonal changes in head height/carapace length ratio. The pattern of seasonal changes of body proportions was similar in all size classes and isometric growth of the head was reported for D. cucullata. Unlike the head, tail spine length/carapace length ratio almost did not vary seasonally. Strong negative allometry of tail spine growth was observed. These results are consistent with the hypothesis that helmets and tail spines provide protection against invertebrates in the two smallest, thus most endangered species.     

Seasonal variations in the condition of two Daphnia species and their hybrid in a eutrophic lake: evidence for food limitation

As seston and chlorophyll concentrations in eutrophic lakesare usually high all year round, it was hypothesized in thepast that food limitation is of major importance for the seasonaldynamics of herbivorous zooplankton in such lakes. Since directmeasurements of food are hampered by the lack of knowledge onthe exact nature of the food in eutrophic conditions, indirectmeasurements are necessary to estimate the degree of food limitationin these circumstances. Hence, we used laboratory- and field-derivedrelationships between the body length and body carbon contentof different species of Daphnia, which were collected from thehighly eutrophic Tjeukemeer, the Netherlands. From a seasonalsurvey of the carbon content of the daphnids. we concluded thatin Tjeukemeer D.galeata, D.galeata x cucullata and D.cucullataare food limited during the largest part of the year. Sincethe condition of the hybrids was relatively high as comparedwith the parental species when the food concentration was high,D.galeata x cucutlaia is expected to be the more successfultaxon during periods of high food availability.     

Generational and dual-species exposures to invertebrate predators influence relative head size in Daphnia mendotae

Laboratory cyclomorphosis experiments with crustacean zooplanktonoften fail to produce morphological trait expression as extremeas that observed in field populations. A seven-generation laboratoryexperiment found that the maximum expression of relative headarea in a round-helmet morph of Daphnia mendotae was delayeduntil the third generation of non-lethal, chemical exposureto Leptodora kindtii. By the third generation and after, relativehead area remained high and comparable in degree to field specimens.The induction pattern from parents to offspring was statisticallypositive into the fourth generation. Published studies on kairomoneinduction of morphological trait expression extending only onegeneration may fail to measure transgenerational effects, whichcould account for mismatch between laboratory and field results.In a second laboratory experiment with a pointed-helmet morphof D. mendotae exposed to L. kindtii and Chaoborus punctipennis,we show that induction levels are not the same for all predatorsor combinations of predators. The maximum relative head areafor pointed morph individuals exposed to both predator cueswas still not as extreme in the laboratory as in the field.This is the first demonstration that delayed generational effectsof kairomone induction of helmet expression in the Cladoceracan reconcile laboratory experiments with the field observationsof cyclomorphosis.     

Behavioural and morphological changes in polymorphic Daphnia related to different predation regimes

Morphology and behaviour of Daphnia galeata in Lake Lille Lungegårdsvann, Bergen, Norway, were studied in two consecutive years. A portion of the population showed helmet cyclomorphosis in 1992, but no helmets developed in 1993. The non-helmeted individuals in 1992 were found in the littoral habitat during summer, whereas the helmeted morph stayed mainly in the pelagial. These differences in morphology and behaviour within and between years are likely to be caused by changes in predation regimes. Larvae of the predacious phantom midge, Chaoborus flavicans, were observed only in 1992, whereas the three-spined stickleback, Gasterosteus aculeatus, was present both years.