Many crustaceans have complex life cycles characterized by a metamorphosis, yet variation in metamorphic traits, and the causes and consequences of this variation, have rarely been examined. Food concentrations were changed during specific larval stages of the freshwater copepod Mesocyclops edax Forbes (Copepoda: Cyclopoida) to examine whether age and size at metamorphosis remain flexible or become fixed during the larval period. Results were compared to predictions of both flexible (the Wilbur—Collins model) and fixed (Leips—Travis model) rate models for the timing of amphibian metamorphosis. Age and size at metamorphosis were variable in all treatments, and age was always more variable than size. Changes in food concentration early in larval development resulted in significant differences in age at metamorphosis among treatments, but changes initiated when 60% of the larval period had passed had no effect on age at metamorphosis. Development appeared to become fixed later in the larval period, before the ultimate larval stage was reached. These results support predictions of the Leips—Travis model. Early changes in food concentrations had significant effects on size at metamorphosis, but changes initiated during the penultimate larval stage (50—60% of larval development) had no effect on metamorph size. Size at metamorphosis in M. edax also appeared to be fixed before the ultimate larval stage was reached. Fixation of size at metamorphosis during development is not predicted by either model and may be unique to organisms with rigid exoskeletons that constrain growth within any stage. Patterns of covariation between age and size at metamorphosis suggest that food conditions early in larval development exert a large effect on metamorphic traits, in contrast to patterns observed in several amphibian species. The Wilbur—Collins model places a fitness premium on delaying metamorphosis to achieve a maximum size, when growth conditions are favorable; it thus may not apply to crustaceans. Selection pressures on the timing of metamorphosis in crustaceans may differ substantially from those identified for amphibians and other organisms. Because of these differences, incorporating crustaceans into studies of metamorphosis will help to clarify the factors affecting this life cycle transition.

Variation in the abundance of animals has traditionally been explained as the outcome of endogenous forcing from density dependence and exogenous forcing arising from variation in weather and predation. Emerging evidence suggests that the effects of density dependence interact with external influences on population dynamics. In particular, spatial heterogeneity in resources and the presence of capable predators may weaken feedbacks from density dependence to growth of populations. We used the Kalman filter to analyze 23 time series of estimates of abundance of northern ungulate populations arrayed along a latitudinal gradient (latitude range of 40°-70°N) to evaluate the influence of spatial heterogeneity in resources and predation on density dependence. We also used contingency tables to test whether density dependence was independent of the presence of carnivores (our estimate of predation) and multiple regressions to determine the effects of spatial heterogeneity in resources, predation, and latitude on the strength of density dependence. Our results showed that the strength of density dependence of ungulate populations was low in the presence of large carnivores, particularly at northern latitudes with low primary productivity. We found that heterogeneity in elevation, which we assume acted as a surrogate for spatial heterogeneity in plant phenology, also reduced effects of density dependence. Thus, we show that external forces created by heterogeneity in resources and predation interact with internal feedbacks from population density to shape dynamics of populations of northern ungulates.

JSTOR is a not-for-profit service that helps scholars, researchers, and students discover, use, and build upon a wide range of content in a trusted digital archive. We use information technology and tools to increase productivity and facilitate new forms of scholarship. For more information about JSTOR, please contact support@jstor.org.. Ecological Society of America is collaborating with JSTOR to digitize, preserve and extend access to Ecology.Abstract. Food quality often has profound effects on life history traits and individual fitness, altering rates of growth and development, changing the timing of reproduction, and shifting the trade-off between egg size and egg number. Few data are available on the effects of food quality on copepod life history traits. We measured several life history traits on a large number of individuals to document the effects of food quality on individual traits, on life history correlates, and on a composite measure of individual fitness in the freshwater copepod Boeckella triarticulata Thomson. Nauplii were raised individually on two diets: one consisted of the high quality alga Cryptomonas sp. (abbreviated as CR), and the second diet consisted of a combination of Cryptomonas sp. and the low quality cyanobacterium Anabaena flos-aquae (CA). The mixed CA diet slowed growth and development so that individuals raised on this diet were older and smaller at metamorphosis and maturity. Despite these effects, there were no differences between diets in survival to maturity, and male copepods raised on the mixed diet lived significantly longer than females or than either sex raised on Cryptomonas alone. Females raised on the mixed diet produced more and larger clutches than those raised on CR, so that total egg production increased on this diet, although large intradiet variation obscured statistical differences between diets in these parameters. Intradiet variation was due to a large range in the number of clutches produced by individual females: some individuals produced 10-15 clutches, contradicting previous descriptions of this species as semelparous. Although diet affected age at first reproduction, it had no significant effect on individual fitness, estimated as X. Boeckella triarticulata achieved high fitness either by minimizing age at first reproduction (CR diet) or by increasing reproductive output (CA diet). Data collected from cast-off exuviae allow nondestructive measures of individual life history traits along with a composite measure of individual fitness. Combining these two analyses is an important step in unraveling life history correlates and in identifying the selective forces driving life history evolution in these crustaceans.

Methods for estimating zooplankton instar development rates and survivorships from cohort analyses are often based on two equations that are correct only under limited and ecologically unrealistic conditions. We derive more general forms of the two equations and show that, using them, development times and survivorships cannot both be calculated from the information available in a single cohort analysis.