Evolution of life histories and yields in experimental populations of Daphnia magna

An experiment on life-history evolution is described in which replicated populations of the Cladoceran Daphnia magna , made up of a standard mixture of clones, were subjected to two contrasting culling regimes, involving removal of small or large individuals. After approximately 150 days of culling, analysis of the life histories showed that genetic differences had emerged between culling regimes. Clones selected by culling small sized individuals grew rapidly through small size classes, whereas those selected by culling large sizes grew slowly through small size classes, with the result that the age at which they became vulnerable to harvesting was delayed. In addition, there was some redistribution of reproduction towards size classes that were not culled. This evolution is consistent with a major decline observed in the yield from populations in which large individuals were culled. We argue that changes of this general kind are likely to take place in exploited populations and that serious consideration should be given to evolutionary aspects of the management of such populations.     

The interaction of Chaoborus size and vertical distribution determines predation effects on Daphnia

1. Larvae of Chaoborus, the phantom midge, are important pelagic planktivores in many freshwater lakes and ponds. The effect of Chaoborus on its prey depends on its size, especially mouth gape diameter, and vertical migration pattern, which affects predator–prey spatial overlap. These two features vary considerably in different Chaoborus species and instars. In this study, the interacting effects of both Chaoborus size and vertical distribution on population growth of Daphnia pulex was analysed with a field enclosure experiment and a matrix population model. 2. In the field experiment, Daphnia were grown in four replicated treatments that included a control (no Chaoborus) and three combinations of instar III and IV Chaoborus of two species (C. trivittatus and C. americanus). Parameters of the matrix model were based on differences between Chaoborus species and instars in capture and ingestion of Daphnia of differing sizes (prey vulnerability) and in vertical overlap with Daphnia in each treatment (density risk). 3. In comparison with the control, the two treatments containing the smaller, migratory C. americanus showed a significant effect on Daphnia population growth rate, while the treatment containing only the larger, non‐migratory C. trivittatus did not. The model accurately simulated these effects. 4. A Daphnia predation risk model, which uses prey vulnerability and density risk parameters, illustrated the individual and combined effect of the different Chaoborus types on Daphnia. Daphnia have a high prey vulnerability to the large C. trivittatus, but overall predation risk was low because of very little overlap. On the contrary, the smaller C. americanus affects only a small range of Daphnia instars, each with a low vulnerability, yet those instars that were vulnerable had a very high density risk because of an increased overlap. 5. This analysis of Daphnia predation risk parameters with coexisting Chaoborus species strongly supports an integrated approach using both size and vertical distribution to determine the ultimate predation effect on Daphnia.     

The response of female guppies, Poecilia reticulata, to chemical stimuli from injured conspecifics

Predation is an important component of the life history and behavior of many fish. In some species of teleost fishes, conspecifics respond to chemical stimuli from injured conspecifics by displaying a fright response, such as increased cohesion of schooling. The purpose of this experiment was to investigate whether female guppies, Poecilia reticulata, display a fright response when exposed to a filtered extract made from wounded conspecifics. The schooling behavior of female guppies was observed for ten minute periods for each of three treatments: baseline treatment (no substances added to the aquarium water), sham treatment (distilled water added to the aquarium water), and alarm treatment (fish extract added to the aquarium water). Female guppies responded with a fright reaction when exposed to a filtered extract prepared from wounded conspecifics; female swam in a more cohesive school during the alarm treatment than they did in either the baseline or sham treatments. The guppy fright reaction response indicates that the fish extract is recognized by conspecifics. This revised version was published online in August 2006 with corrections to the Cover Date.     

Relative importance of inefficient feeding and consumer excretion to organic carbon flux from Daphnia

1. Sloppy and inefficient feeding by zooplankton is generally thought to make a major contribution to the regeneration of the dissolved organic carbon (DOC) pool in aquatic environments. In this study, we tested experimentally the regeneration of DOC by a freshwater zooplankter feeding on two species of phytoplankton at different food concentrations and C : P ratios. We separated the DOC production because of inefficient feeding (pre‐ingestive regeneration) and zooplankton excretion and faeces release (postdigestive regeneration). 2. Within a brief incubation period (10 min), DOC production in the presence of Daphnia was not significantly different from that in the control treatment without grazers. During a longer incubation period (4 h), the amounts of radiocarbon retained in the algal cells per se were constant or were not different from those in the control treatments. These experimental results strongly suggest that inefficient feeding did not contribute significantly to DOC production in the grazer–prey system. 3. During the 4‐h incubation, calculations of the DOC per ingestion rate (i.e. DOC produced by Daphnia alone) showed that food concentration and algal species did not affect the relative DOC production, but there was considerable difference at different algal C : P ratios and grazer densities. We found that direct excretion of DOC by Daphnia occurred rapidly following food digestion and accounted for >65% of the total DOC production. Maximum DOC leakage from Daphnia faeces contributed less to DOC production than the grazer excretion, except under P‐limited conditions. 4. This study highlights the dominant role of postingestive process, especially the direct excretion by zooplankton, in DOC production in a grazer–prey system.     

微颗粒紫菜粉投喂量对隆线溞（ Daphnia carinata） 种群增长的影响

为了探索微颗粒紫菜粉在枝角类中培养的效果,采用Banta液、微颗粒紫菜粉（0．12mg／mL、0．16mg／mL和O．20mg／mL）、紫菜粉酶解液（1．2％、1．6％和2．0％）3种食物投喂隆线潘（Daphniacarinata）,并通过隆线潘的种群密度变化,比较不同食物及投喂量下的培养效果。结果表明：以紫菜粉酶解液组培养效果最好,尤以1．6％酶解液组,种群密度最高,达5．73ind／mL,且高峰维持时间较长;投喂微颗粒紫菜粉的培养效果不及酶解液组,以O．16my,／mL组培养效果最好,最高密度为2．87ind／mL;Banta液培养组的潘群密度较稳定,种群最高密度为3．33ind／mL,但培养的潘体较小。实验还显示,培养液的pH值以8．0～8．2时培养效果较佳。     

Costs, benefits and the evolution of inducible defences: a case study with Daphnia pulex

Phenotypic plasticity is one major source of variation in natural populations. Inducible defences, which can be considered threshold traits, are a form of plasticity that generates ecological and evolutionary consequences. A simple cost-benefit model underpins the maintenance and evolution of these threshold, inducible traits. In this model, a rank-order switch in expected fitness, defined by costs and benefits of induction between defended and undefended morphs, predicts the risk level at which individuals should induce defences. Here, taking predator-induced morphological defences in Daphnia pulex as a threshold trait, we provide the first comprehensive investigation into the costs and benefits of a threshold trait, and how they combine to reflect fitness and predict the switchpoint at which induction should occur. We develop reaction norms that show genetic variation in switchpoints. Further experiments show that induction can confer a survival benefit and a cost in terms of lifetime reproductive success. Together, these two traits combine to estimate expected fitness and can predict the switchpoint between an undefended and a defended strategy. The predictions match the reaction norm data for clones that experience these costs and benefits, and correspond well to independent field data on induction. However, predictions do not, and cannot, match for clones that do not gain a benefit from induction. This study confirms that a simple theory, based on life history costs and benefits, is a sufficient framework for understanding the ecology and evolution of inducible, threshold traits.     

Variation in the neonate size of Daphnia pulex: the effects of predator exposure and clonal origin

Daphnia pulex clones originating from twelve small pond habitats were exposed to chemical cues from a size-selective predator, larvae of the phantom midge Chaoborus. Exposure delayed the onset of reproduction and increased the size at first reproduction. On the other hand, the neonates produced by these larger mothers were slightly smaller than the neonates produced by the smaller control mothers. In cladocerans, neonate size is usually positively correlated to the size of the mother. Thus exposure to Chaoborus kairomone apparently had direct effects on neonate size counterbalancing the maternal effects. Daphnia clones collected from Chaoborus-free and Chaoborus-rich environments exhibited different responses. In first adult instar, the clones from Chaoborus inhabited environments increased their offspring size under Chaoborus exposure whereas clones from Chaoborus-free environments did not. This may reflect clonal adaptation to the predation prehistory of their original habitat since larger neonates more quickly reach a size protected from the predator.     

Finding the optimal vertical distribution: behavioural responses of Daphnia pulicaria to gradients of environmental factors and the presence of fish

1. The vertical distribution of zooplankton results from active habitat choice aiming to optimise fitness gain in a system of trade‐offs. 2. Using large, controlled indoor mesocosms (Plön Plankton Towers), we monitored the behavioural response of Daphnia pulicaria to vertical gradients of temperature, food, oxygen and light, in the presence and absence of fish predation. 3. In the absence of fish, Daphnia distributed as predicted by an ideal ‘free distribution with costs’. If the food was distributed homogeneously, they stayed in the warm epilimnion, while they balanced their time dwelling in epi‐ and hypolimnion if the food was concentrated in a deep‐water maximum. 4. However, oxygen depletion in the hypolimnion, representing an additional cost, prevented Daphnia from completely exploiting the hypolimnetic food maximum. Consequently, the proportion dwelling in the hypolimnion was larger if oxygen was not limiting. 5. Fish predation had an overwhelming effect, driving Daphnia into the hypolimnion under all experimental conditions. If permitted by oxygen availability, Daphnia used the whole hypolimnion, but oxygen depletion reduced their possible habitat to the upper hypolimnion with oxygen concentrations above c. 0.7 mg L^?1. As fish were less tolerant of low oxygen, the layer below the thermocline formed a predation refuge for Daphnia.     

Colonization dynamics and life history traits of seven Daphnia pulex genotypes

Summary A series of experiments revealed significant differences in the potential ability of seven Daphnia pulex genotypes to colonize two lakes in which this species does not naturally reside. Life table experiments, in which individuals from each genotype were raised separately on water and natural phytoplankton from the two lakes, revealed several significant differences among genotypes in life history traits, including age and size at first reproduction, clutch size and offspring body size. Significant differences among genotypes were also found in mean genotype fitness and rate of population increase, although all genotypes were able to increase in absolute numbers. Significant genotypexlake interaction was found for several life history traits and mean fitness, indicating that the relative success of invading genotypes may depend on habitat characteristics. Enclosure experiments, in which all seven genotypes were introduced together into enclosures in both lakes, revealed that some genotypes increased greatly while others declined in relative abundance, and that the most successful genotypes differed between lakes. In addition, the most successful genotypes in the enclosures were not necessarily the genotypes that displayed the highest fitness in the life table experiments, possibly because individuals in the enclosures competed for food resources, leading to exclusion of certain genotypes.     

Effects of salinity on metabolism and life history characteristics of Daphnia magna

In the Baltic Sea area, the cladoceran Daphnia magna is commonly found in brackish water rockpools and it has been suggested that salinity is one of the niche dimensions that affects the distribution of the species. The salinity tolerance of D. magna was studied both in physiological and life history experiments. The experimental salinities were freshwater, 4S and 8S. The highest respiration and ammonium excretion rates were measured in the freshwater treatment with decreasing respiration and ammonium excretion rates at higher salinities. The lowest O/N ratio (oxygen consumption to ammonium excretion), describing the metabolic status of an organism, was obtained at 8S, although the only significant differences were detected when comparing to 4S treatments. Individual growth rate, reproductive output and population growth rate were highest at 4S. At 8S growth and reproduction were reduced as compared to freshwater and 4S. The life history parameters in the performed experiments indicated higher fitness (expressed as r) as well as more favourable conditions for growth and reproduction at 4S, whereas the O/N ratio was more difficult to interpret and, in this case, gave a less clear picture of the salinity influence.     

Effect of food conditions on the bacterial feeding of Daphnia galeata

Feeding selectivities of Daphnia galeata were measured using various density combinations of ¹⁴C-labeled Chlamydomonas and ³H-labeled natural bacteria. In all food combinations, D. galeata preferred algae. Selectivity coefficient (bacterial clearance rate: algal clearance rate) was 0.35 at the level of incipient food limitation (1.5 × 10⁶ µm³ ml^–1).However, it increased up to 0.9 with increasing total food concentration. A simple mechanical sieving model alone cannot explain such a change in the selectivity coefficient. A mechanism of clogging suggestively explain the increase of bacterial ingestion efficiency by D. galeata, especially at high food concentrations. The change of selectivity coefficient related to food condition suggests that bacterial feeding by the Daphnia is incidental event associated with the algal feeding.     

Differing effects of suspended sediments on the performance of native and exotic Daphnia

1. Although large cladocerans are usually uncommon in large rivers, Daphnia lumholtzi (an exotic species in North America) is widespread and occasionally abundant. We collected zooplankton on the Illinois River (Illinois, U.S.A.) in 1995 and 1996 and found that the peak density of D. lumholtzi (25 L⁻¹) typically exceeded that of all native species combined. Maximum density occurred during warm periods (up to 27 °C) when concentrations of inorganic suspended sediments were high (>50 mg L⁻¹).
2. Using a life table experiment, we examined the effects of variation in suspended sediment (0 and 80 mg L⁻¹) and food (10⁴ and 10⁵ Ankistrodesmus cells mL⁻¹) on fitness of D. lumholtzi and the native Daphnia parvula. Daphnia lumholtzi had greater survivorship than D. parvula in most treatments and higher life-time fertility than D. parvula in all treatments. Both species achieved their fastest intrinsic rates of growth in treatments with high food, but their responses to suspended solids differed. The growth rate of D. lumholtzi in high food was slightly increased by higher turbidity, whereas that of D. parvula was depressed.
3. Results suggest that the ability of D. lumholtzi to tolerate suspended solids is an important factor contributing to its success in invading North American rivers.     

Patterns in the life histories and feeding strategies of British macrolepidoptera

Temperate Lepidoptera typically have life cycles of a year or less. The time available to feed and attain a suitable state for reproduction might thus constrain potential life-history patterns, and species characteristics such as adult size, overwintering stage, first month of larval feeding and feeding specificity may only occur in particular combinations. Several studies have documented correlates of feeding ecology which suggest this is indeed the case. We use data for more than 900 British macrolepidoptera to establish relationships among several ecological and life-history variables to determine whether only particular combinations can occur. Few patterns of feeding characteristics predicted in the literature are observed. Distinct, consistent life-history syndromes cannot be defined; almost any combination of the life-history characteristics we looked at can occur. So long as it is possible to fit in with timing constraints at certain critical times of year, there seems to be a great deal of flexibility in how the rest of the time can be filled. However, several broad and potentially interesting patterns still emerge. Polyphagous species are larger than specialists; the month in which species start to feed inlinked to both their overwintering stage and the growth form of the plant they feed upon; and there are no species overwintering in the egg stage and having two broods per year.     

Life histories and habitat selection in Daphnia: divergent life histories of D. magna clones differing in phototactic behaviour

To test the hypothesis of co-adaptation of life histories and daytime vertical distribution (vertical migration behaviour) in Daphnia, life history characteristics were analysed for two positively, three negatively, and four intermediately phototactic Daphnia magna clones. Clones with different phototactic behaviour were found to have divergent life history strategies, with positively phototactic clones being good exploiters under the non-limiting conditions provided in the laboratory, i.e. low density (1 ind./1), high food concentration (6,5–7 10⁵ Scenedesmus cells/ml, restored daily) and high temperature (20° C). They realized a high intrinsic rate of increase at a small adult body size through rapid development, at a cost of producing small neonates. Negatively and intermediately phototactic clones had larger adult body sizes, and produced larger neonates that were more starvation-resistant than those of positively phototactic clones. Selection for high intrinsic rate of increase in intermediately phototactic clones was mediated through the production of large clutches.     

Seasonal variation of egg size and number in a Daphnia pulex population

Seasonal variation of egg size and number was examined in a Daphnia pulex population inhabiting a vernal pond. In this population, size at maturity declines at midseason, probably as an adaptive response to size-selective predation by larvae of the salamander Ambystoma. The larger early season individuals produce more and larger eggs than the smaller late season individuals. Age at maturity does not vary between seasons. Laboratory experiments indicate that temperature may affect egg size, egg number and size at maturity. However, field data suggest that temperature accounts for only a small fraction of the total variation in egg size and number. Indirect measures of nutrition indicate that food limitation does not cause the seasonal decline in egg size and number. The seasonal change in reproductive traits is well correlated with changes in invertebrate and vertebrate predation. Examination of predator feeding preferences and their impact on Daphnia mortality indicate that variation of reproductive traits is most likely a complex adaptation to changing predation regimes.     

Influence of predation by fish and water turbidity on a Daphnia gessneri population in an Amazonian floodplain lake,Brazil

The population behavior of Daphnia gessneri Herbst, 1967 in a floodplain lake (Lago Grande) of the lower Rio Solimões was investigated between April 1979 and March 1980 with regard to 1) predation by the fish called tambaqui (Colossoma macropomum, Characidae), 2) water level fluctuation and 3) water transparency. Zooplankton density samples were collected at two sites near mid-lake, where water depth and Secchi disc transparency were measured. In addition, qualitative samples of zooplankton and fish collections were taken at several sites in the adjacent floodplain areas. The author concludes that fluctuations in Daphnia gessneri populations correlate most with intense predation by fish and water turbidity.     

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.     

Specificity of predator-induced neck spine and alteration in life history traits in Daphnia pulex

It has been proposed that the predator-induced defensive neck spine in Daphnia pulex has a demographic cost. Our results show that this cost is not merely an allocation cost related to the formation and maintenance of the neck spine. In a life table experiment, we tested whether spine induction and life history traits in D. pulex are affected by different invertebrate predators: first and third instar Chaoborus, fourth instar Mochlonyx and two size classes of Notonecta and Dytiscus larvae. D. pulex showed sensitivity to the different predators. Predator-exposure affected one or more of the following life history traits of D. pulex: the timing of first reproduction, clutch size, and growth. In some cases, exposure to predators altered life history traits when neck spine induction did not occur. These shifts in life history traits occurring in the absence of spine induction may be caused by behavioral or physiological changes triggered by the predators.     

A laboratory study of the feeding of Mesostoma lingua (Schmidt) (Turbellaria: Neorhabdocoela) on Daphnia magna Straus at four different temperatures

Mesostoma lingua eats 0.8 Daphnia magna at 15 °C, and 2.04 at 30 °C. Below 0.5 Daphnia per day (D d^–1) hunger occurs, at any temperature, and mortality increases sharply. The functional response curve appears to be linear, but saturation was not fully reached at 5 D d^–1, the highest food level applied in the present study. All food regimes (above hunger level) and temperature regimes differed significantly from each other; factor interactions were of increasing importance at higher temperatures.Longevity, above hunger level, decreased with temperature. Peak longevity at low temperature may be weakly associated with relatively low food levels.     

Effects of salinity on the life history and fitness of Daphnia magna: variability within and between populations

The life history traits of Daphnia magna were studied in laboratory experiments under freshwater and brackish (5 salinity) conditions. The variability of responses within and between populations was examined by comparing 11 clones from a brackish lake and 10 clones from a freshwater pond. Experimental clones were hatched from ephippia collected from the sediment and thus represent random samples of the clone banks of each population.Most clones with a high salinity tolerance were from the population of the brackish habitat, but some were also found in the freshwater population. Thus, freshwater populations appear to have the potential to invade brackish habitats. A proportion of clones from the brackish population had very low fitness (measured as e^r) under freshwater conditions. This unexpected result means that freshwater adaptation can be lost by the freshwater cladoceran Daphnia magna. The effects of unfavourable conditions on growth and reproduction varied among clones and were not correlated. This clonal variation in growth and reproduction indicates that the environmental sensitivities of these traits are independent. The pattern of fitness reaction norms showed no trade-off between fitness under brackish and under freshwater conditions for either population. Thus, euryhaline generalists should be favoured in habitats with salinity fluctuations between freshwater and brackish conditions.