Retarded growth of cladoceran zooplankton in the presence of a copepod predator

Various instars of four different cladoceran species representing a wide spectrum of body size were grown at high food availability in the presence and in the absence of natural densities of an invertebrate predator, a cyclopoid copepod Acanthocyclops robustus (G.O. Sars). Daily weight increments calculated from individual weights at the end and at the beginning of each 1, 2 or 4 day experiment, showed that individual growth was more or less drastically retarded in the presence of the predator as well as when exposed to water in which the predator had been feeding. The data also showed that the effect of this invertebrate predator was more pronounced in small prey instars and small prey species that were more vulnerable to predation than large prey.     

Interacting effects of predation risk and food availability on larval anuran behaviour and development

Age and size at metamorphosis are two important fitness components in species with complex life cycles. In anurans, metamorphic traits show remarkable phenotypic plasticity, especially in response to changes in growth conditions. It is also possible that the perception of risk directly determines changes in larval period and the size of metamorphs. This study examines how the perception of predation risk affects the timing of and size at metamorphosis in common frogs (Rana temporaria). I raised tadpoles at two risk levels (fish-conditioned water or unconditioned water) crossed with the availability or lack of food at night (all tadpoles had food available in the day). Tadpoles reacted to chemical cues from predatory fish by decreasing activity. A novel behavioural result was a predation×food interaction effect on refuge use, which also accounted for most of the predator main effect: predation risk only caused increased refuge use in the night-starved treatment. Despite these behavioural modifications, the perception of predation risk did not affect growth rate and mass at metamorphosis in a simple way: the effects of food regime on growth and size at metamorphosis were dependent on the level of predation risk as revealed by significant predation×food interaction effects. Tadpoles who had food withheld at night metamorphosed at the smallest size, suggesting a negative relationship between size at metamorphosis and refuge use. Tadpoles raised in fish-conditioned water had longer larval periods than those in unconditioned water, but these differences were significant only if food was available at night. These results conflict with the hypotheses that tadpoles should reduce their larval period or growth rates (and hence metamorphose at a smaller size) as the risk of predation increases. In contrast to predation risk, food availability strongly affected the length of the larval period: night-starved tadpoles metamorphosed relatively early with or without fish stimulus. Thus, early metamorphosis resulted from periods of low food availability, but not from a heightened ”perceived risk” of predation. This example counters the hypothesis of acceleration of the developmental rate (which shortens the time to metamorphosis) as a mechanism to escape a risky environment. Received: 18 August 1999 / Accepted: 10 January 2000     

Interacting effects of predation risk and resource level on escape speed of amphibian larvae along a latitudinal gradient

Fast‐growing genotypes living in time‐constrained environments are often more prone to predation, suggesting that growth‐predation risk trade‐offs are important factors maintaining variation in growth along climatic gradients. However, the mechanisms underlying how fast growth increases predation‐mediated mortality are not well understood. Here, we investigated if slow‐growing, low‐latitude individuals have faster escape swimming speed than fast‐growing high‐latitude individuals using common frog (Rana temporaria) tadpoles from eight populations collected along a 1500 km latitudinal gradient. We measured escape speed in terms of burst and endurance speeds in tadpoles raised in the laboratory at two food levels and in the presence and absence of a predator (Aeshna dragonfly larvae). We did not find any latitudinal trend in escape speed performance. In low food treatments, burst speed was higher in tadpoles reared with predators but did not differ between high‐food treatments. Endurance speed, on the contrary, was lower in high‐food tadpoles reared with predators and did not differ between treatments at low food levels. Tadpoles reared with predators showed inducible morphology (increased relative body size and tail depth), which had positive effects on speed endurance at low but not at high food levels. Burst speed was positively affected by tail length and tail muscle size in the absence of predators. Our results suggest that escape speed does not trade‐off with fast growth along the latitudinal gradient in R. temporaria tadpoles. Instead, escape speed is a plastic trait and strongly influenced by the interaction between resource level and predation risk.     

ADAPTIVE GENETIC VARIATION IN GROWTH AND DEVELOPMENT OF TADPOLES OF THE HYBRIDOGENETIC RANA ESCULENTA COMPLEX

The distribution and proportion of the sexual species Rana lessonae to the hemiclonal hybrid R. esculenta among natural habitats suggests that these anurans may differ in adaptive abilities. I used a half-sib design to partition phenotypic and quantitative genetic variation in tadpole responses at two food levels into causal variance components. Rana lessonae displays strong phenotypic variation across food levels. Growth rate is strictly determined by environmental factors and includes weak maternal effects. Larval period and body size at metamorphosis both contain moderate levels of additive genetic variance. The sire x food interactions and the lack of environmental correlations indicate that adaptive phenotypic plasticity is present in both of these traits. In contrast, R. esculenta displays less phenotypic variation across food levels, especially for larval period. Variation in body size at metamorphosis is underlain by genetic variation as shown by high levels of additive genetic variance, yet growth rate and larval period are not. Significant environmental correlations between larval period at high food level and growth, larval period, and body size at low food, indicate phenotypic plasticity is absent. A positive phenotypic correlation between body size at metamorphosis and larval period for R. lessonae at both food levels suggests a trade-off between growing large and metamorphosing quickly to escape predation or pond drying. The lack of a similar correlation for R. esculenta at the high food level suggests it may be less constrained. Different levels of adaptive genetic variation among larval traits suggest that the sexual species and the hybridogenetic hemiclone differ in their abilities to cope with temporally and spatially heterogeneous environments.     

Developmental response of the brown-winged green bug, Plautia stali (Heteroptera: Pentatomidae), to food shortage

To study developmental response of the brown-winged green bug, Plautia stali Scott, to food shortage we reared nymphs under restricted feeding conditions produced by shortening the feeding period after molt or withholding food from second to fifth instars. For second instars, molting rates were significantly reduced as the feeding period was shortened. Shortening the feeding periods for third to fifth instars also reduced molting rates, but less so; some nymphs were able to complete their developments even if no food was given to any of the instars. Compared with controls for which feeding was not restricted, nymphs that successfully reached the next instar had reduced postmolt body size as a result of restricted feeding for all instars except the fifth (2-day feeding for second instar, and 1-day feeding for third and fourth instars), whereas instar duration was unchanged or only slightly prolonged for all instars. These results suggest that only nymphs with nutritional accumulation over a specific threshold in each instar can progress to the next instar, and that, particularly for second to fourth instars, nymphs develop on schedule without prolonging instar durations to compensate for reduced growth under conditions of food shortage.     

The effect of host developmental stage at parasitism on sex‐related size differentiation in a larval endoparasitoid

• 1 For their larval development, parasitoids depend on the quality and quantity of resources provided by a single host. Therefore, a close relationship is predicted between the size of the host at parasitism and the size of the emerging adult wasp. This relationship is less clear for koinobiont than for idiobiont parasitoids.
• 2 As size differentiation in host species exhibiting sexual size dimorphism (SSD) is likely to occur already during larval development, in koinobiont larval endoparasitoids the size of the emerging adult may also be constrained based on the sex of the host caterpillar.
• 3 Sex‐specific growth trajectories were compared in unparasitised Plutella xylostella caterpillars and in second and fourth instar hosts that were parasitised by the solitary larval koinobiont endoparasitoid Diadegma semiclausum. Both species exhibit SSD, where females are significantly larger than males.
• 4 Healthy female P. xylostella caterpillars developed significantly faster than their male conspecifics. Host regulation induced by D. semiclausum parasitism depended on the instar attacked. Parasitism in second‐instar caterpillars reduced growth compared to healthy unparasitised caterpillars, whereas parasitism in fourth‐instar caterpillars arrested development. The reduction in growth was most pronounced in hosts producing male D. semiclausum.
• 5 Parasitism itself had the largest impact on host growth. SSD in the parasitoid is mainly the result of differences in growth rate of the parasitoid–host complex producing male and female wasps and differences in exploitation of the host resources. Female wasps converted host biomass more efficiently into adult biomass than males.

     

Clutch size and body size at first reproduction in Daphnia pulicaria at different levels of food and predation

Field data from seven alpine lakes in Serra da Estrela. Portugal.show that reproduction in Daphnia may be as efficiently controlledby fish predation and copepod predation on eggs in brood cavitiesas it is by food limitation. Body length and clutch size estimatesin Daphnia pulicaria revealed high inter- and intra-populationvariability in maturation size (body size at first reproduction).and in number of eggs per clutch. Daphnia at first maturationin lakes stocked with rainbow trout were half the size of thosefound in fishless lakes (body length of 0.86–0.95 and1.55–1.81 mm. respectively). The mean number of eggs perclutch was reduced to a similar degree by food limitation, predationby fish and copepod predation on eggs in brood cavities, butthe underlying mechanisms of this reduction were different.Food limitation caused smaller clutch sizes in all individuals,so variation remained the same. Fish predation caused the selectiveremoval of individuals with maximum clutches, so variation decreased.Copepod predation caused removal of eggs from brood cavitiesof randomly infested females, so that variation increased, particularlyat a high food level when non-infested females carried largeclutches of eggs.     

Reproductive allometry does not explain the temperature-size rule in water striders (Aquarius remigis)

Most ectotherms follow the temperature-size rule meaning that individuals growing up under cool conditions are larger as adults than those growing up in warm conditions. This pattern is difficult to explain because growth is usually slower in the cold meaning it takes longer to reach a larger adult size. One potential explanation for this pattern is that the typical increase in fecundity with body size is steeper in cool environments than it is in warm environments. As such, the relative gain in fecundity for being larger in the cold would compensate for the extra time it takes to grow to that size. We present the first empirical test of this model using the water strider Aquarius remigis. Individuals were reared at either 20° or 25°C with subsampling at each instar to estimate growth trajectories. At adult eclosion, half were switched to the alternate temperature and all females isolated, mated, and reproductive output measured for 3 weeks. We found that A. remigis does follow the temperature-size rule but that fecundity was highly plastic with respect to laying temperature such that the slightly greater fecundity of those reared at 20°C was due to their larger size. Overall, those laying at 25°C were more fecund and showed a positive relationship between body size and fecundity. Those laying at 20°C did not show a significant relationship between size and fecundity. As such, the reproductive allometry shows a pattern reverse to what would be needed for a larger size in the cold to be adaptive in this species. Although A. remigis follows the temperature-size rule over this temperature range, this is likely due to a constraint on growth and development rather than being adaptive plasticity.     

Predation on the invasive mosquito Aedes japonicus (Diptera: Culicidae) by native copepod species in Germany

Some limnic copepod species are predators of mosquito larvae. Seven species belonging to the order Cyclopoida, family Cyclopidae, were collected in the field in Germany and tested for the first time in laboratory bioassays for their potential to serve as biological control agents of the invasive Asian bush mosquito Aedes japonicus (Theobald), a vector of various pathogens causing disease. Females of Diacyclops bicuspidatus (Claus) did not attack 1^st instar larvae of Ae. japonicus, but Macrocyclops distinctus (Richard), Cyclops divergens Lindberg, and C. heberti Einsle predated a mean of 14, 18, or 19 1^st instar larvae, respectively. Acanthocyclops einslei Mirabdullayev and Defaye killed 30 larvae, and high predation rates with a mean of 39 or 46 larvae, respectively, were obtained by Megacyclops viridis (Jurine) and M. gigas (Claus). In regression analyses, predation rates by M. viridis correlated with body size, with specimens of 1.8 mm length being more effective than smaller or bigger ones. Based on the presented data, the two Megacyclops species seem to be promising candidates for use in field studies on the biological control of Ae. japonicus.     

Food and growth relations of the marine microzooplankter,Synchaeta cecilia (Rotifera)

The trophic interactions of the marine rotifer Synchaeta cecilia were investigated by determining its feeding and growth rates on a wide variety of marine phytoplankton and by determining its susceptibility to predation by the calanoid copepod, Acartia tonsa. Reproduction of S. cecilia was sustained in four-day feeding trails by 13 of 37 algal species tested. Growth-supporting species included species of Cryptophyceae, Dinophyceae, Chlorophyceae and Haptophyceae in sizes from 4 to 47 μm. Within these taxa, other species in the acceptable size range failed to support growth. No species of Cyanophyceae, Bacillariophyceae, or Chrysophyceae supported growth of the rotifer. S. cecilia can be maintained on unialgal cultures of Cryptophyceae but growth is enhanced by a combination of two or three species; a mixture of Chroomonas salina (Cryptophyceae), Heterocapsa pygmaea (Dinophyceae), and Isochrysis galbana (Haptophyceae) has sustained laboratory stocks of S. cecilia for over four years. The expected response of S. cecilia to food quantity was observed: as food concentration was increased from 58 to 1154 μg C 1⁻¹, the population growth constant increased from 0.17 to 0.60 d⁻¹ at 20°C. This is equivalent to population doubling times of 4.0 and 1.1 days at H. pygmaea densities of 500 and 10⁴ cells ml⁻¹, respectively. The susceptibility of S. cecilia to predation was investigated by determining its rate of capture by the omnivorous marine copepod Acartia tonsa. At prey densities of 5 to 35 μg C 1⁻¹ (0.3 to 1.9 individuals 1⁻¹), A. tonsa readily ingested S. cecilia at rates up to 3 μg C copepod⁻¹ day⁻¹.     

Absence of predation eliminates coexistence: experience from the fish–zooplankton interface

Examples from fishless aquatic habitats show that competition among zooplankton for resources instigates rapid exclusion of competitively inferior species in the absence of fish predation, and leads to resource monopolization by the superior competitor. This may be a single species or a few clones with large body size: a cladoceran such as Daphnia pulicaria, or a branchiopod such as Artemia franciscana, each building its population to a density far higher than those found in habitats with fish. The example of zooplankton from two different fish-free habitats demonstrates the overpowering force of fish predation by highlighting the consequences of its absence. Released from the mortality caused by predation, a population of a superior competitor remains at a density equal to the carrying capacity of its habitat, in a steady state with its food resources, consisting of small green flagellate algae, which are successful in compensating high loss rates due to grazing, by fast growth. In such a situation, the high filtering rate of Daphnia or Artemia reduces resources to levels that are sufficient for assimilation to cover the costs of respiration (threshold food concentration) in adults but not in juveniles. This implies long periods of persistence of adults refraining from producing live young, because production of instantly hatching eggs would be maladaptive. Severe competition for limiting resources imposes a strong selective pressure for postponing reproduction or for producing resting eggs until food levels have increased. Offspring can only survive when born in a short time window between such an increase in food levels and its subsequent decline resulting from population growth and intense grazing by juveniles. Such zooplanktons become not only a single-species community, but also form a single cohort with a long-lifespan population. The observations support the notion that diversity may be sustained only where predation keeps densities of coexisting species at levels much below the carrying capacity, as suggested by Hutchinson 50 years ago.     

Interactive effects of temperature and concentration of the flavonol rutin on growth, molt and food utilization of Manduca sexta caterpillars

A factorial experiment tested the effects of varying concentrations of the flavonol rutin and daytime temperatures of 20 and 30°C on growth, molting and food utilization efficiencies of third instar tobacco hornworms (Manduca sexta (L.)). Cool temperature prolonged both the growth (=feeding) and non-feeding periods and consequently the relative consumption rates (RCR) and relative growth rates (RGR). Temperature had no impact on the amount of food consumed and the utilization indices of efficiency of conversion of ingested food and efficiency of conversion of digested food to larval biomass. But rutin was more concentrated in the frass of the larvae at the warm temperature. With increasing levels of rutin in the diet, the efficiency of conversion of ingested food tended to decline. Rutin reduced RCR and RGR. At the cool temperature, rutin increased the time spent in the first portion of the non-feeding period disproportionately. Analysis of growth rate intervals within the growth period indicated that at the cool temperature rutin had no discernible impact over the first half of the growth period, during which developmental competence to molt is likely achieved. Overall, these results indicate an overlap in the growth and molting phases and suggest that rutin interferes with physiological processes at the time of molt initiation, with these effects magnified at a cool temperature.     

EFFECTS OF BT RICE ON THE FOOD CONSUMPTION,GROWTH AND SURVIVAL OF CHILO SUPPRESSAUS LARVAE UNDER DIFFERENT TEMPERATURES*

Abstract The Bt transgenic rice line, i.e. KMD1 containing a synthetic cry 1 Ab gene from Bacillus thuringiensis Berliner was tested in the laboratory for evaluating its effects on larval food consumption, growth and survival of striped stem borer (SSB), Chilo suppressalis (Walker) under different temperatures. The food consumption and body weight growth as well as the survival of both 3rd and 5th instar larvae fed on Bt rice were significantly reduced, as compared with those on the control. The temperature showed no marked effect on both food consumption and body weight growth of larvae fed on Bt rice stems since the onset of 3rd instar, conversely had significant effects as since the initial stage of 5th instar. In contrast, the temperature did not distinctly influence the survival of both 3rd and 5th instar larvae. There was a linearly positive relationship between the corrected mortality of larvae and the accumulative amount of Bt rice tissues ingested by larvae.     

Juvenile survival of a planktonic insect: effects of food limitation and predation

• 1 An experiment was conducted to investigate potential impacts of food limitation and copepod predation on juvenile survival of Chaoborus purtctipennis. We tested the hypotheses that: (i) juvenile survival of Chaoborus is influenced more by copepod predation than by starvation in a productive environment, and (ii) food limitation and predation interact to affect survival.
• 2 Effects of food concentration (approximately 800, 1400 and 2300 microzooplankton 1-^?1) and predator density (0, 1 and 2 Mesocydops edax 1-^?1) on Chaoborus development and survival were evaluated using a 3 × 3 factorial design. Jars containing lake water, the appropriate food and predator treatments, and two Chaoborus (<12h old) were rotated on a plankton wheel at 25°C. Survival and developmental stage were monitored daily until all individuals had either died or moulted to instar II.
• 3 Predation by Mesocydops was the major source of mortality, causing 87.5% of Chaoborus deaths over all treatments. Chaoborus mortality was significantly higher in treatments with Mesocydops (67–100%) than in predator-free treatment (0–13%).
• 4 Development time was significantly longer in the low-density food treatment than in the highest food treatment.
• 5 No significant interaction between food limitation and predation was detected.
• 6 These results suggest that predation by copepods may limit recruitment of juvenile Chaoborus in productive lakes.

     

Regulation of rotifer densities by crustacean zooplankton in an oligotrophic montane lake in British Columbia

Summary To examine the relative demographic effects of predation and competition for food in rotifers during spring and summer in an oligotrophic lake, predator and competitor densities and food supplies were experimentally altered inside large enclosures. Abundances of rotifer species were positively correlated with experimental densities of fourth instar Chaoborus trivittatus larvae, a major crustacean predator in 1976. Experimental alteration of the densities of Daphnia rosea and Diaptomus leptopus and D. kenai in 1978 produced highly significant increases in rotifer biomass only under Daphnia removal, but not under copepod removal. Inorganic fertilizer additions to enclosures in 1978 and 1979 revealed minimal rotifer increases unless pulsed additions were large or Daphnia were also excluded. Large demographic reponses of rotifers to low fertilizer loadings in the absence of Daphnia confirmed the pre-eminence of competitive food limitation in producing rotifer scarcity in summer.     

Preliminary results of an experimental study of the effects of mysid predation on estuarine zooplankton community structure

In a shallow estuarine system near Beaufort, North Carolina, a period of high winter abundance of the mysidsMysidopsis bigelowi andNeomysis americana was associated with a change in zooplankton species composition, from dominance byAcartia tonsa to dominance byCentropages spp. andSaphirella sp. Both mysids feed onA. tonsa at higher rates than the other copepods. Experiments were carried out in 600–1 000 liter enclosures, in which the initial mysid density was manipulated and the effects on the enclosed copepod community were monitored. Mysid predation had a significant effect on copepod densities. The effects of mysid predation on species composition appeared to depend on the relationship between their prey preferences and the dominant copepod species present in the communities. Under conditions favoring dominance byA. tonsa, the preferred prey species, the results suggested that mysid predation may reduce dominance and increase diversity. But when the less preferredCentropages was dominant, mysid predation had no effect on species composition.     

Egg-brooding, body size and predation risk in planktonic marine copepods

We investigated the interacting effects of copepod body size and the presence or absence of egg masses on the risk of predation by a visual predator. We conducted selection experiments involving three-spined sticklebacks (Gasterosteus aculeatus) and copepods ranging in body mass from 0.5 to 740 μg C: Oithona similis, Corycaeus anglicus, Pseudocalanus newmani, P. moultoni, Pseudodiaptomus marinus, and Paraeuchaeta elongata. We found that sticklebacks selected ovigerous females of the two smallest-bodied species of copepods (Oithona similis and Corycaeus anglicus). In contrast, the fish showed no significant selection for ovigerous females of the remaining, larger-bodied species. Unexpectedly, egg mass position (i.e., in a ventral, dorsal or lateral location on the urosome) appeared to influence predation risk more than did body size, resulting in higher predation risk for the cyclopoid and poecilostomatoid species than for the calanoid species we tested. Although the sticklebacks showed no statistically significant preference for ovigerous females of any of the four calanoid species, for each species the overall proportion of ovigerous females ingested was slightly greater than 0.50. Thus, whether body size influences the susceptibility of egg-brooding calanoid copepods to predation remains an open question. Received: 24 August 1998 / Accepted: 6 July 1999     

Temperature-mediated changes in zooplankton body size: large scale temporal and spatial analysis

Climate warming has been linked with changes in the spatiotemporal distribution of species and the body size structure of ecological communities. Body size is a master trait underlying a host of physiological, ecological and evolutionary processes. However, the relative importance of environmental drivers and life history strategies on community body size structure across large spatial and temporal scales is poorly understood. We used detailed data of 83 copepod species, monitored over a 57-year period across the North Atlantic, to test how sea surface temperature, thermal and day length seasonality relate to observed latitudinal-size clines of the zooplankton community. The genus Calanus includes dominant taxa in the North Atlantic that overwinter at ocean depth. Thus we compared the copepod community size structure with and without Calanus species, to partition the influence of this life history strategy. The mean community body size of copepods was positively associated with latitude and negatively associated with temperature, suggesting that these communities follow Bergmann's rule. Including Calanus species strengthens these relationships due to their larger than average body sizes and high seasonal abundances, indicating that the latitudinal-size cline may be adaptive. We suggest that seasonal food availability prevents high abundance of smaller-sized copepods at higher latitudes, and that active vertical migration of dominant pelagic species can increase their survival rate over the resource-poor seasons. These findings improve our understanding of the impacts that climate warming has on ecological communities, with potential consequences for trophic interactions and biogeochemical processes that are well known to be size dependent.     

Antennule shape and body size of <Emphasis Type="Italic">Bosmina</Emphasis>: key factors determining its vulnerability to predacious Copepoda

Cyclopoid copepods are common in lakes and ponds, and they have a significant predation impact on the communities of the small zooplankton species. To reduce the predation risk, some cladoceran zooplankters develop protuberant (defensive) morphologies in the presence of the copepods. In the case of the small cladoceran Bosmina, they elongate their appendages (antennule and mucrone) and change the antennule morphotype. However, information about the effectiveness of these defensive devices against copepod predation is still insufficient. In our study, to find the compositive effects of these appendages on the vulnerability of Bosmina, we exposed two bosminid species (B. longirostris and B. fatalis) of different body sizes and with appendages of different lengths and shapes to copepod (Mesocyclops) predation. The experiment revealed that the shape of the antennule is a main factor determining the bosminid’s vulnerability to copepod predation and indicated that the protection of the opened ventral carapace must be a key strategy by which Bosmina avoids copepod predation.     

Resource limitation,predation risk and compensatory growth in a damselfly

Periods of poor nutrition during early development may have negative fitness consequences in subsequent periods of ontogeny. In insects, suppression of growth and developmental rate during the larval stage are likely to affect size and timing of maturity, which in turn may lead to reduced reproductive success or survivorship. In light of these costs, individuals may achieve compensatory growth via behavioural or physiological mechanisms following food limitation. In this study, we examined the effects of a temporary period of food restriction on subsequent growth and age and size at maturity in the larval damselfly Ischnura verticalis (Odonata: Coenagrionidae). We also asked whether this temporary period of reduced nutrition affected subsequent foraging behaviour under predation risk. I. verticalis larvae exposed to a temporary food shortage suffered from a reduced growth rate during this period relative to a control group that was fed ad libitum. However, increased growth rates later in development ensured that adult body size measurements (head and pronotum widths) did not differ between the treatments upon emergence. In contrast, adult dry mass did not catch up to that of the controls, indicating that the increased growth rates for size dimensions occur at the cost of similar gains in mass. Predators reduced foraging effort of larvae, but this reduction did not differ between control larvae and those previously exposed to poor nutrition.