The synergistic effects of temperature and food concentration of life history parameters of Daphnia

Summary Daphnia parvula experiences a wide range of thermal (4°–30°C) and food conditions during its annual population cycle. We used cohort life table experiments to determine the synergistic effects of temperature and food concentration on its life history parameters. A 3×3 factorial design experiments was conducted with animals raised at all combinations of three naturally experienced temperatures (10°, 15° and 25° C) and food levels (0.02, 0.20 and 2.0 mg C/l). D. parvula showed an increase in survivorship with decreasing temperature at all food levels. Fecundity parameters (number of broods/female, brood size and net reproductive rate) increased with increasing food at the two lower temperatures but showed a mid-range food optimum at the highest temperature. Development rates and realized rates of increase (r) showed an increase with both increasing temperature and food such that they were maximum at the highest temperature-food level treatment. The life history parameters, average lifespan, age at first reproduction, brood duration time, brood size and number of young per reproductive female all showed significant interaction between temperature and food as was suggested by trends in R _o and r. Temperature had a reduced effect on fecundity, development rates and realized rates of increase at the low food level.Population birth rates of continuously reproducing zooplankton are typically calculated by the egg-ratio method (Edmondson 1960) and are based on thermally controlled development rates which neglect the effects of food limitation. Significant synergistic temperature-food effects on brood duration time and other life history parameters of Daphnia parvula suggest that food limitation and foodtemperature interaction should be considered when calculating field population birth rates. A comparison of realized rates of increase from this study with similar life table data for Daphnia parvula raised on natural food from Lake Oglethorpe indicate that synergistic effects are negligible in this eutrophic system due to abundant resources. However, synergistic effects are probably important in oligotrophic systems where resources are limited.This work was supported by National Science Foundation Grant DEB 8005582. Lake Oglethorpe Limnological Association, Contribution No. 24 and Harbor Branch Foundation, Inc. Contribution No. 369     

Life history of Daphnia galeata in a hypertrophic reservoir and consequences of non-consumptive mortality for the initiation of a midsummer decline

SUMMARY 1. Field and laboratory investigations were combined in a 2‐year study on the initiation of a midsummer decline of Daphnia galeata Sars in a hypertrophic reservoir. Quantitative field samples were taken twice a week, and, adult and juvenile mortality rates were calculated. Patterns of reproduction and survival of daphnids born during spring and early summer under fluctuating food conditions were determined in life‐table experiments. 2. The abundance of Daphnia increased strongly in early May and declined in June 1998 (midsummer decline). In 1999, Daphnia density increased only slowly in spring and remained constantly high throughout the summer. 3. Food conditions (concentrations of POC_{<30 μm}) for daphnids deteriorated in both years in response to increasing Daphnia densities, resulting in a clear‐water phase of about 4 weeks. When Daphnia abundance declined in 1998, POC_{<30 μm} concentrations increased greatly, whereas in 1999 food conditions improved only slightly and Secchi depth remained high. 4. Survival of daphnids in life‐table experiments decreased greatly after food became rare and was strongly reduced in those animals born during the clear‐water phase compared with those born later. In addition, age at first reproduction was retarded during the clear‐water phase, resulting in very low population growth rates. Survivorship patterns in life‐table experiments suggest a strong impact of non‐consumptive mortality on Daphnia population dynamics. 5. Field data of mortality point to differences in mortality patterns between years, probably resulting from different predation impacts of juvenile fish. In both years, however, adult mortality contributed substantially to overall mortality at the end of the clear‐water phase. As bottom‐up effects on D. galeata were very similar in both years, the significance of non‐consumptive mortality on the initiation of midsummer declines appears to depend largely on recruitment patterns before the clear‐water phase. A high impact can be expected when Daphnia populations are dominated by a peak cohort of nearly identical age during the clear‐water phase.     

Photoperiod and temperature as inducers of gamogenesis in a dicyclic population of Scapholeberis armata Herrick (Crustacea: Cladocera: Daphniidae)

The induction of gamogenesis in a dicyclic population of Scapholeberis armata from a small pond at 40° N in Pennsylvania, USA, was studied. Animals were collected at the time of gamogenesis in mid-October (fall) 1988 and early June (spring) 1989). 10 clones were established from isolated females for each season and maintained at low density under standard conditions of 20 °C and 13.5L/10.5D, with Chlamydomonas reinhardtii diluted with pond water to 3 · 105 cells ml^–1 as a food source. For each clone, mothers of pairs of test animals were acclimated to 4 sets of conditions: 20 °C/15.5L (spring), 15 °C/11.5L (fall), 20 °C/11.5L (control), and 15 °C/15.5L. Test animals were observed bidaily or daily for up to 37 days. Occurrence of molts and of parthenogenetic broods or ephippia was recorded. At short photoperiod, ephippial females developed, usually after one or more parthenogenetic broods, among both spring and fall animals. The response was stronger (higher % sexual females) among fall clones, and was enhanced in both groups by 15 °C water. No animals responded to long photoperiod in warm or cool water. It is likely that additional factors, such as prior photoinduction, rapid depletion of food supply, and increased inter-animal contact due to high population density are required for the induction of spring gamogenesis. Differences in the responsiveness of spring and fall clones to inducing conditions suggest some differences in genotypes of spring and fall subpopulations in this pond. It is proposed that variable responses among genotypes to factors inducing gamogenesis may be the basis for the differences and clines in the timing of gamogenesis found among ecologically and latitudinally separated populations.     

Biotic relations affecting species structure in zooplankton communities

Mathematical modelling and laboratory experiments were used to study how exploitative competition and predation influence the species structure in cladoceran community. For five species of Cladocera (Sida crystallina, Daphnia magna, Simocephalus vetulus, Daphnia longispina, and Diaphanosoma brachyurum), representing a gradient of body size, population characteristics were described as functions of food concentration. Abundance dynamics were simulated in mixed species cultures and invasion experiments under different levels of food supply corresponding to oligo-, meso-, and eutrophic conditions. Separate simulations were also run including and excluding (fish) predation. The competitive ability of each species was estimated as the values of the population equilibrium food concentration. Simulation results showed that for the no-predator scenario, increases in the level of food supply promoted species coexistence while under lower food concentrations only one species remained at the end of the simulation runs. When predation was allowed, the number of species that coexisted at the end of the simulations increased up to four species, indicating that predator pressure facilitated species coexistence because it shortened periods of food depletion. Simulation results were verified in laboratory experiments which suggested that population equilibrium food concentration can be used as an estimate of competitive ability. Finally, species structure and relative abundance in Lake Naroch (Belarus) during the summer of 2004 was found to be consistent with our results from computer simulation and laboratory experiments with regard to competition and predation impacts on zooplankton community. Handling editor: S. I. Dodson     

The role of food quality in clonal succession in Daphnia: an experimental test

A high genetic variation and recurrent changes in the genetic structure have been found in many pelagic populations. However, evidence that directly links these changes to differences in the ecological performance of particular genotypes is scarce. We hypothesized that within Daphnia, the specialization of clones occurring in a particular season to the food quality specific for that time of the year is responsible for the observed changes in the genetic structure of a population. This hypothesis was tested by comparing the fitness of spring and summer clones of the Daphnia longispina group, given food of biochemical quality relevant to these seasons. We identified significant intraspecific differences between clones of Daphnia that are specific for particular seasons, but there was no evidence that clones are adapted to the food quality available at the respective times of year. Summer clones reproduce at smaller size, and have a lower juvenile specific growth rate as compared to spring clones, irrespective of food quality. Spring clones invest more energy in somatic growth at the cost of reproduction, whereas summer clones invest more energy in reproduction at the cost of somatic growth. On the basis of the observed differences between spring and summer clones in their patterns of energy allocation, we suggest that other factors, most likely predation, are the major forces driving phenotypic and genetic diversity in the investigated Daphnia population of a large lake.     

Life history variation among low-arctic clones of obligately parthenogenetic Daphnia pulex: a diploid-polyploid complex

Summary Laboratory life table experiments were conducted using nine clones of obligately parthenogenetic Daphnia pulex that were collected from a site in the Canadian low-arctic. Two of the nine clones were diploids, while the other seven clones were polyploids. Significant clonal differences in age at first reproduction, size at first reproduction, number of offspring in each of the first three broods, offsrring sizes for the first two broods, and intrinsic rates of natural increase were detected. Differences in life histories were evident between polyploids and diploids. Generally, polyploid clones reached maturity at later ages, matured at larger sizes, produced smaller broods, and larger offspring than the diploid clones. The data are discussed in reference to potential biotic (i.e. invertebrate predation) and abiotic factors (i.e. physicochemical gradients) that may influence life history variation in this clonal assemblage.     

Significance of a low oxygen layer for a Daphnia population in Lake Yunoko,Japan

Population dynamics and vertical migration of Daphnia longispina in Lake Yunoko were studied. The Daphnia population was small in spring and early summer, probably because of high predation pressure by fish. The population grew in midsummer, when thermal stratification developed and the dissolved oxygen became very low in the deeper layer of the hypolimnion. In this season, adults of D. longispina concentrated in the daytime near the lake bottom, where fish were absent because of the anoxic conditions, but ascended at night to the upper layer of the hypolimnion, where food was most abundant. The low oxygen layer near the bottom kept out the predators and protected Daphnia from predation, and consequently contributed to the built-up of its population. However, the low oxygen layer was unfavorable for reproduction of Daphnia, as reflected in the low egg ratio and high percentage of males in the population. The population decreased in the fall, when thermal stratification disappeared and predation pressure seemed to increase.     

Temperature and kairomone induced life history plasticity in coexisting Daphnia

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

Dietary restriction in two rotifer species: the effect of the length of food deprivation on life span and reproduction

According to resource allocation theory, animals face a trade off between the allocation of resources into reproduction and into individual growth/maintenance. This trade off is reinforced when food conditions decline. It is well established in biological research that many animals increase their life span when food is in suboptimal supply for growth and/or reproduction. Such a situation of reduced food availability is called dietary restriction. An increase in life span under dietary restricted conditions is seen as a strategy to tolerate periods of food shortage so that the animals can start reproduction again when food is in greater supply. In this study, the effect of dietary restriction on life span and reproduction in two rotifer species, Cephalodella sp. and Elosa worallii, was investigated using life table experiments. The food concentration under dietary restricted conditions was below the threshold for population growth. It was (1) tested whether the rotifers start reproduction again after food replenishment, and (2) estimated whether the time scale of dietary restricted conditions is relevant for the persistence of a population in the field. Only E. worallii responded to dietary restriction with an increase in life span at the expense of reproduction. After replenishment of food, E. worallii started to reproduce again within 1 day. With an increase in the duration of dietary restricted conditions of up to 15 days, which is longer than the median life span of E. worallii under food saturation, the life span increased and the life time reproduction decreased. These results suggest that in a temporally (or spatially) variable environment, some rotifer populations can persist even during long periods of severe food deprivation.     

Life history comparisons of Daphnia obtusa from temporary ponds, cultured with a low-quality food

Temporary pond Daphnia flourish on relatively poor-quality food, suggesting adaptation to stringent temporary pond conditions. We conducted laboratory life history experiments on populations of Daphnia obtusa from a shaded woodland temporary pond (short hydroperiod, dystrophic) and an open farmland temporary pond (long hydroperiod, eutrophic), and compared a suite of physical/chemical conditions in the ponds to evaluate habitat conditions. We hypothesized that the shaded woodland pond population would be more fit in terms of life history variables for individuals (age and size at first reproduction, mean brood size, mean number of neonates) and populations (generation time, net reproductive rate R ₀, and intrinsic reproductive rate r) given a standard, low-quality food (trout chow and yeast). Life history traits of woodland pond animals were mixed, relative to farm pond animals, and consistent with bet-hedging for an unpredictable habitat. Values of life history traits rivaled or exceeded those of other studies using phytoplankton as food, and were influenced by the pond water used for our study. Life histories clearly differ among local and regional temporary pond Daphnia obtusa populations, and should be valuable for examining the relative influences of local selection and metapopulation dynamics on population structure.     

Absence of typical diel vertical migration in Daphnia: varying role of water clarity, food, and dissolved oxygen in Lake Eymir, Turkey

Diel vertical migration (DVM) is a complex and dynamic behaviour against predation because the reaction of migrating organisms to light intensity plays a primary role, but is modified by other factors. In the relatively shallow but thermally stratified Lake Eymir, Daphnia pulex de Geers utilized vertical refugia afforded by the hypolimnion during both day and night. Differences in general vulnerability to fish predation determined the differences in their mean residence depths (MRDs) of different population categories such as most conspicuous and vulnerable individuals of adult with eggs inhabited the deepest depth, whereas juveniles stayed close the thermocline. In late spring, profoundly high amplitude of displacement within the hypolimnion, probably due to the hypolimnion being well-lit and relatively well-oxygenated for the fish and rather unsafe for the large-sized daphnids, was recorded. Therefore, the large-sized daphnids daytime refuge was close to the bottom whereas at night they moved upward to benefit from warmer water temperature along with food availability in the presence of fish predation but still remained below the thermocline. In summer, the insignificant amplitude of the hypolimnetic, which later became epilimnetic, displacements were probably due to the near-anoxic condition found below the thermocline. This might have deterred the fish, thus providing a safer refuge for daphnids in the below thermocline, which afterwards became the above thermocline. Low oxygen availability was regarded as the summer proximate factor. The abundant food and warmer water conditions found in the below/above thermocline also accounted for absence of DVM in summer. Consequently, this study suggests that DVM by Daphnia is an adaptation that is plastic to changing environmental conditions.     

Effects of food limitation,notonectid predation,and temperature on the population dynamics of Daphnia carinata

The relative contributions of invertebrate predation (Notonectidae: Anisops spp.), food limitation, and certain abiotic factors in driving the population dynamics of Daphnia carinata were studied for a two year period in a large farm dam in southern Victoria, Australia. Detailed measurements were made on the population densities of Daphnia and Anisops spp., the amount of food available, the nutritional status, and the size-specific fecundity of Daphnia. The density of the Daphnia population at the field site oscillated closely with water temperature. The amplitude of the population fluctuations varied seasonally, being much greater during the warmer months of the year and switching to fluctuations with low peaks when water temperature dropped below approximately 15°C. Anisops spp. densities were greatest in winter and declined during the spring of each year. Nymphs appeared in late spring and early summer and numerically dominated the population. During the warmer periods of the year, the daphnid population went through a series of rapid growth phases leading to over-exploitation of food resources and subsequent population collapses. Daphnia population densities were not correlated with Anisops numbers suggesting that predation was not a major regulatory factor during the warmer periods of the year. When water temperatures fell below ca. 15°C daphnid population densities remained low despite high food levels. During this period the impact of Anisops may have been greater. Two distinct phases were identified: a warm water period when food limitation was the main regulatory factor, and a cool water period when Anisops predation may have been the paramount factor. Low oxygen concentrations were associated with heavy rainfall in the spring and may have had a limiting effect on Daphnia for short periods. Daphnia may have had an important role in sustaining the Anisops population in the pond over each winter.     

Life history, growth and production of Mysis relicta in the large, fiord-type Lake Mjøsa, Norway

SUMMARY.

• 1 Biomass, production and life history of Mysis relicta were studied in the large Lake Mjøsa during the years 1976–80. Biomass fluctuations were large, but biomass averaged about 1 g wet wt m^?2 or 200 ind. m^?2. Cumulative net production during summer ranged from 1.6 to 2.1 g wet wt m^?2. Mean population P/B-ratio was close to 2.
• 2 Neither production or biomass of Mysis were correlated with the biomass of phytoplankton or the main Zooplankton prey species (Daphnia and Bosmina), suggesting a predominant ‘top down’ control of Mysis in this lake.
• 3 The life cycle was well synchronized with development of food items, but predation pressure also affected life-history patterns. The proportion of juveniles apparently increased in years with strong fish predation pressure, producing a predominantly 1-year life-cycle pattern the following year.
• 4 Mysis had a predominant 2-year life history, but a traction of the population may reproduce after 1 or 3 years. Reproduction occurred exclusively during winter. Even in such a large and stable system, where Mysis has coexisted with its main food items and predators for 8–10 000 years, a flexible life history is maintained and is probably an important buffer against year-to-year fluctuations in food and predator abundance.

     

Effects of Food Availability and Predation on a Marine Zooplankton Community — a Study on Copepods in the Baltic Sea

The importance of food resources versus predatory effects in determining the abundance pattern of zooplankton varies over time and space. Here, we evaluate the population dynamics of dominant calanoid copepods (Acartia spp. bifilosa/A. longiremis and Eurytemora affinis) in a reference area and a eutrophied area in the northern Baltic Sea Proper. We base the paper on a combination of statistical analyses of a long term (12 years) data set on population biomass dynamics, laboratory experiments on the feeding biology of these species and literature data of their susceptibility to predation. Overall, copepods were equally abundant at both sites, but Eurytemora was more abundant in the eutrophied area while Acartia spp. dominated in the non-eutrophied reference area. There was, however, no significant difference between the two areas in biomass dynamics (biomass specific rate of change) of the two groups of copepods. However, in spring and early summer the population biomass increased faster for Eurytemora as compared to Acartia spp. This is consistent with our findings from feeding and respiration experiments. Eurytemora exhibited generally higher ingestion rates and had a higher growth efficiency (ratio of ingestion to respiration rates), which suggests an advantage over Acartia spp. when food availability increases. Both species generally showed a rapid population decline starting in late summer, most likely caused by intensive predation by fish and mysid shrimps. In late autumn and winter, biomasses of both genera continued to decrease, but the decline was fastest for Eurytemora. During this period of the year, when the food availability is sparse, Acartia spp. may have an advantage over Eurytemora, as we found that the former ingests a broader size range of food particles and it has also been reported to be able to switch from filter feeding to raptorial feeding. This study shows that dominant copepods in the net-zooplankton community in our study area were structured both by food availability and predation. The relative importance of these effects varied seasonally.     

Impact of larval Dorosoma predation on Daphnia parvula dynamics

• 1 In reservoirs of the south-eastern United States, Daphnia typically decline in abundance during the summer months. Current understanding suggests that adult shad (Dorosoma spp.), the dominant planktivore in these reservoirs, plays a limited role in Daphnia midsummer declines. The impact of shad larvae is beginning to be documented.
• 2 To examine the impact of larval shad on Daphnia seasonal dynamics, larval dynamics were studied over two seasons and stomach contents were analysed during one season.
• 3 Initial Daphnia panwla declines correlated with abundance of shad larvae. During the decline, shad stomach contents consisted of 97—100%D. parvula. After D. parvula declined, Bosmina and Diaphanosoma accounted for up to 97% of larval diets. When shad diets included other cladocerans, D. parvula abundance increased in the reservoir.
• 4 When this predation pattern was incorporated into a predictive model, larval shad predation accounted for an initial, temporary decline in D. parvula abundance. However, when other crustaceans were incorporated into the shad diet, D. parvula populations increased.
• 5 These results suggest that larval shad have a temporary, strong influence on the structure and abundance of D. parvula populations in south-eastern reservoirs of the United States.

     

A comparison of life‐history and parental care in temperate and tropical wrens

Comparing closely related species that live in different environments is a powerful way to understand selective pressures that influence life‐history evolution. We examined a suite of life‐history traits and parental care in neotropical buff‐breasted wrens Cantorchilus leucotis and north‐temperate Carolina wrens Thryothorus ludovicianus (Family Troglodytidae), to test hypotheses about life‐history evolution. As expected, buff‐breasted wrens exhibited smaller clutch sizes and higher annual adult survival than Carolina wrens. We found minimal support for the nest predation hypothesis, as nest survival and age‐corrected provisioning rates to whole broods were similar between species, and number of breeding attempts and breeding season length were greater in temperate wrens. Critical predictions of the food limitation hypothesis were not supported; in particular age‐corrected provisioning rates per nestling were higher in the tropical than temperate species. The adult survival and offspring quality hypothesis garnered the most support, as buff‐breasted wrens exhibited greater age‐corrected provisioning rates per nestling, a longer nestling period, longer re‐nesting intervals following nest success, and lower annual fecundity than Carolina wrens. Despite similarly prolonged breeding seasons, reproductive strategies differ between species with buff‐breasted wrens investing considerably in single broods to optimize first‐year survival and Carolina wrens investing in multiple small broods to optimize annual fecundity.     

Carry-over effects of the larval environment on post-metamorphic performance in two hylid frogs

Life history theory and empirical studies suggest that large size or earlier metamorphosis are suitable proxies for increased lifetime fitness. Thus, across a gradient of larval habitat quality, individuals with similar phenotypes for these traits should exhibit similar post-metamorphic performance. Here we examine this paradigm by testing for differences in post-metamorphic growth and survival independent of metamorphic size in a temperate (spring peeper, Pseudacris crucifer) and tropical (red-eyed treefrog, Agalychnis callidryas) anuran reared under differing larval conditions. For spring peepers, increased food in the larval environment increased post-metamorphic growth efficiency more than predicted by metamorphic phenotype and led to increased mass. Similarly, red-eyed treefrogs reared at low larval density ended the experiment at a higher mass than predicted by metamorphic phenotype. These results show that larval environments can have delayed effects not captured by examining only metamorphic phenotype. These delayed effects for the larval environment link larval and juvenile life history stages and could be important in the population dynamics of organisms with complex life cycles.     

Cannibalistic behaviour of rock-pool copepods: An experimental approach for space, food and kinship

Studies on cannibalism in harpacticoid copepods are restricted to predation on naupliar larvae in rock-pool harpacticoids of the genus Tigriopus. An earlier experimental study on the Mediterranean copepod Tigriopus fulvus indicated that females recognized their own larvae and preferentially preyed on nauplii other than their own. In a series of laboratory experiments, we tested if there were differences in naupliar predation as a function of crowding, food level and sex in Tigriopus brevicornis and T. fulvus. Results show that cannibalism was restricted to the first larval stages (N1 and N2). Both food availability and adult density significantly affected the predation rate. Contrary to earlier suggestions, adult males also preyed on the nauplii. We found no evidence that adults spare their own offspring, for neither T. fulvus nor T. brevicornis.This is in accordance with what one would expect for species having the life history characteristics of Tigriopus, i.e.: multiple broods and large number of offspring. Earlier results indicating parental care in Tigriopus must be taken with caution.     

Investigations on fecundity of Bythotrephes longimanus in Lake Lucerne (Switzerland) and on Niche Segregation of Leptodora kindti and Bythotrephes longimanus in Swiss lakes

In Lake Lucerne, Switzerland, the predaceous cladocerans Leptodora kindti and Bythotrephes longimanus segregate along spatial and temporal dimensions. In spring (April–May/June), Bythotrephes longimanus occurs below 0–20 m, while Leptodora is absent. In summer and early autumn (July–September/October), when Leptodora dominates during daytime in the 0–20 m depth, Bythotrephes longimanus also lives in deeper zones. Food competition and fish predation pressure may be the cause of differences in ecology of Leptodora and Bythotrephes acquired during evolution. Due to its transparency and tolerance of higher temperature, Leptodora could avoid fish predation and, therefore, competes with Bythotrephes longimanus successfully. In addition, the differences between the two species may account for the spatial and temporal niche segregation in oligotrophic Swiss Lakes. But spatial niche segregation is less important in mesotrophic lakes with high prey density than in oligotrophic lakes with low prey density. In small, eutrophic lakes importance of temporal niche segregation also decreases, and Bythotrephes is seldom or not present. The preference of Bythotrephes to live in deeper water to avoid fish predation during summer may be the cause of its difficulties to establish itself in small and eutrophic lakes with high prey densities, where the hypolimnion is missing or anoxic.In the spring, Bythotrephes exhibits r-strategy (smaller body size and a higher fecundity), the female is already fertile after the first molt. In the summer, a K-strategy prevails (larger body length and lower fecundity than in the spring), and female Bythotrephes are fertile only after the second molt. Shortage of prey (biomass of Bosmina and Daphniadecreased after June especially in the surface layers) and the maximum fish predation pressure in summer may change the life strategy of Bythotrephes: while fecundity decreases from generation to generation, body length increases. Enhanced prey densities (e.g. during mesotrophic conditions in L. Lucerne) lead to larger individuals in summer and autumn.     

Warming erodes individual-level variability in life history responses to predation risk in larvae of the mayfly Cloeon dipterum

1. Warming and predation risk are ubiquitous environmental factors that can modify life histories and population dynamics of aquatic ectotherms. While separate responses to each of these factors are well understood, their joint effects on individual life histories and population dynamics remain largely unexplored. Current theory predicts that the magnitude of prey behavioural, physiological, and life history responses to predation risk should diminish with warming due to the reduced metabolic scope. However, empirical support for this prediction remains equivocal, and experiments covering a substantial proportion of individual prey ontogeny until maturation are lacking.
2. To fill these gaps, we ran a laboratory experiment to investigate how warming and non-consumptive predation risk influence life history responses in the larvae of the mayfly Cloeon dipterum, an aquatic insect with highly plastic development. We reared larvae of varying initial sizes at three temperatures (21, 24, and 27°C) in a risk-free environment and under predation risk signalled by chemical cues from dragonfly larvae (Aeshna cyanea), and followed their individual survival, growth, and development until emergence.
3. Some C. dipterum larvae substantially prolonged their development and the proportion of these slow individuals declined rapidly with temperature and increased with predation risk. We attribute this response to cohort splitting, a common life history strategy of aquatic insects and other taxa in unpredictable environment.
4. Growth, development, and maturation varied predictably with temperature in the fast larvae that did not prolong their development. They grew and developed faster but matured at smaller sizes with increasing temperature. Predation risk tended to slow down individual growth and development in line with the reduced metabolic scope hypothesis, but the differences were relatively minor and observable only at 21°C.
5. Survival to subimago increased with predation risk, possibly due to indirect effects mediated by dissolved micronutrients, but did not vary significantly with temperature. Survival also tended to be higher in the slow individuals. This partly compensated for a smaller final size relative to the fast individuals and made both strategies comparable in overall fitness.
6. Our results show that warming may erode individual-level variability in life history responses to predation risk. This implies that warming can synchronise population dynamics and consequently make such populations more vulnerable to unpredictable disturbances.