Tom E. X. Miller

Effects of Different Resource Additions of Species Diversity in an Annual Plant Community

A commonly observed phenomenon in plant communities is that the ad- dition of a limiting resource leads to an increase in productivity and a decrease in species diversity. We tested the hypothesis that the mechanism underlying this pattern is a dis- proportionate increase in mortality of smaller or shade-intolerant species in more produc- tive sites caused by reduction of light levels. We added water and/or one of three nutrients (nitrogen, phosphorus, and potassium) to a 1 st-yr old-field community dominated by weedy annuals and measured effects on productivity, species composition, diversity, and light levels after one growing season. Diversity was not clearly related to productivity in this experiment. Watering increased productivity, but, contrary to expectations, had no effect on density of surviving plants, species diversity, or abundance of low-growing species. Almost all the increase in biomass with watering was due to a positive response by Ambrosia artemisiifolia, an upright annual that was the most common species in the canopy in all treatments. The addition of nitrogen had only a small positive effect on productivity, but strongly decreased density of surviving plants, species diversity, and abundance of most low-growing species. Only Ambrosia increased in abundance with nitrogen addition. The phosphorus and potassium additions had little effect on the community. We suggest that the high mortality and low diversity in the nitrogen addition plots, but not in the more productive watered plots, was due to limitation by nitrogen earlier than limitation by water during the growing season. The consequence was earlier canopy closure and greater mortality due to light limitation.

RESOURCE AND TOP-PREDATOR REGULATION IN THE PITCHER PLANT (SARRACENIA PURPUREA) INQUILINE COMMUNITY

The effects of top-down and bottom-up forces on species abundance and diversity were quantified in the inquiline communities found in the water-filled leaves of the pitcher plant, Sarracenia purpurea. A press field experiment was conducted in which the abundances of resources (dead ants) and the top predators (larva of the mosquito Wyeomiia smithii) were each maintained at three levels in a factorial design for 23 d. Abundances of mites, rotifers, protozoans, and bacteria increased significantly with prey addition. Protozoan species richness also increased with increasing resource levels, whereas additional resources did not affect bacterial species richness. Increases in top-predator density decreased rotifer abundance but increased bacterial abundance and species richness. Increases in bacterial abundance and richness indicate that a trophic cascade occurs via the larvae, rotifer, and bacteria pathway; omnivory appears to be not strong enough to affect the trophic cascade. Top-down and bottom-up forces both had strong, but different, effects on the abundance and richness of intermediate trophic levels. For most components of the inquiline community, species richness and population sizes are resource limited, whereas predator limitation targets a limited set of groups.

Thinking inside the box: community-level consequences of stage-structured populations

Ecologists have historically represented consumer-resource interactions with boxes and arrows. A key assumption of this conceptualization is that all individuals inside a box are functionally equivalent. Demographic stage structure, however, is a widespread source of heterogeneity inside the boxes. Synthesizing recent studies, we show that stage structure can modify the dynamics of consumer-resource communities owing to stage-related shifts in the nature and strength of interactions that occur within and between populations. As a consequence, stage structure can stabilize consumer-resource dynamics, create possibilities for alternative community states, modify conditions for coexistence of competitors, and alter the strength and direction of trophic cascades. Consideration of stage structure can thus lead to outcomes that are not expected based on unstructured approaches.

EFFECT OF COMMUNITY STRUCTURE ON INVASION SUCCESS AND RATE

Although invasion has long been recognized as an important ecological pro- cess, there are very few experimental studies of invasion in natural communities and vir- tually no studies that determine how trophic structure affects the probability of invasion. We introduced novel protozoans and rotifers into the natural communities found in the water-filled leaves of the pitcher plant Sarracenia purpurea. The communities were ma- nipulated in a factorial design of removal of predators (larvae of the mosquito Wyeomyia smithii) and addition of resources (dead insects). Three of the six protozoan species suc- cessfully established populations when introduced into pitchers, suggesting that these spe- cies are migration limited. The other three protozoans and a rotifer did not successfully invade established communities, although all four are naturally found in these inquiline communities. Of the three successfully invading protozoans, two were more likely to invade when resources were added and one of those even more frequently when predators were removed. Invasion by the third was unaffected by these experimental manipulations. Similar effects of predators and resources were found on population sizes of these three species. This study is one of very few that have addressed invasion experimentally; its results suggest that a variety of factors, including migration, predation, and resource availability, can have different influences on invasion by fairly similar protozoans.

Impacts of insect herbivory on cactus population dynamics: experimental demography across an environmental gradient

Understanding the role of consumers in plant population dynamics is important, both conceptually and practically. Yet, while the negative effects of herbivory on plant performance have been well documented, we know much less about how individual-level damage translates to impacts on population growth or whether spatial variation in herbivory affects patterns of plant distribution. We studied the role of insect herbivory in the dynamics and distribution of the tree cholla cactus (Opuntia imbricata), a long-lived perennial plant, across an elevational gradient in central New Mexico, USA, from low-elevation grassland (1670 m) to a grassland–mountain transition zone (1720 m) to the rocky slopes of the Los Pinos Mountains (1790 m). Tree cholla density increased significantly with elevation, while abundance of and damage by a suite of native, cactus-feeding insects decreased. We combined field experiments and demographic models to test the hypothesis that systematic spatial variation in chronic insect herbivory...

Sex-Biased Dispersal and the Speed of Two-Sex Invasions

Population models that combine demography and dispersal are important tools for forecasting the spatial spread of biological invasions. Current models describe the dynamics of only one sex (typically females). Such models cannot account for the sex-related biases in dispersal and mating behavior that are typical of many animal species. In this article, we construct a two-sex integrodifference equation model that overcomes these limitations. We derive an explicit formula for the invasion speed from the model and use it to show that sex-biased dispersal may significantly increase or decrease the invasion speed by skewing the operational sex ratio at the invasion’s low-density leading edge. Which of these possible outcomes occurs depends sensitively on complex interactions among the direction of dispersal bias, the magnitude of bias, and the relative contributions of females and males to local population growth.

Variance in composition of inquiline communities in leaves of Sarracenia purpurea L. on multiple spatial scales

A survey of the abundances of species that inhabit the water-bearing leaves of the pitcher plant Sarracenia purpurea was conducted at several different spatial scales in northern Florida. Individual leaves are hosts to communities of inquiline species, including mosquitoes, midges, mites, copepods, cladocerans, and a diverse bacterial assemblage. Inquiline communities were quantified from four pitchers per plant, three plants per subpopulation, two subpopulations per population, and three populations. Species varied in abundance at different spatial scales. Variation in the abundances of mosquitoes and copepods was not significantly associated with any spatial scale. Midges varied in abundance at the level of populations; one population contained significantly more midges than the other two. Cladocerans varied at the level of the subpopulation, whereas mites varied at the level of the individual plants. Bacterial communities were described by means of Biolog plates, which quantify the types of carbon media used by the bacteria in each pitcher. Bacterial communities were found to vary significantly in composition among individual plants but not among populations or subpopulations. These results suggest that independent factors determining the abundances of individual species are important in determining community patterns in pitcher-plant inquilines.

Evolution of prey in ecological time reduces the effect size of predators in experimental microcosms

Ecologists have long studied the effect of predators on prey population abundance while evolutionary biologists have measured prey trait evolution in response to predation. Ecological and evolutionary processes were generally thought to occur on different time scales, but recent evidence suggests that evolution may alter the ecological effects of predation over the course of ecological experiments. We used a protozoan and its mosquito-larvae predator, naturally found in the water-filled leaves of pitcher plants, to examine the effect of prey evolution on predator-prey interactions. In experiments conducted over 12 days (approximately 50 prey generations, but less than one predator generation), we measured a decrease in the effect of mosquito larvae predators on protozoa prey populations. In a separate set of experiments, we found that the presence of predators corresponded with evolution of smaller cell size and increased population growth rate. In ecological experiments, two situations commonly occur: strong selection pressure applied by the treatment itself and discrepancies in generation times of associate species. Our results suggest that in either situation, the resulting evolutionary patterns may lead to dramatic and important changes in ecological effect size.

Sex and stochasticity affect range expansion of experimental invasions

Understanding and predicting range expansion are key objectives in many basic and applied contexts. Among dioecious organisms, there is strong evidence for sex differences in dispersal, which could alter the sex ratio at the expansions leading edge. However, demographic stochasticity could also affect leading-edge sex ratios, perhaps overwhelming sex-biased dispersal. We used insects in laboratory mesocosms to test the effects of sex-biased dispersal on range expansion, and a simulation model to explore interactive effects of sex-biased dispersal and demographic stochasticity. Sex-biased dispersal created spatial clines in the sex ratio, which influenced offspring production at the front and altered invasion velocity. Increasing female dispersal relative to males accelerated spread, despite the prediction that demographic stochasticity would weaken a signal of sex-biased dispersal. Our results provide the first experimental evidence for an influence of sex-biased dispersal on invasion velocity, highlighting the value of accounting for sex structure in studies of range expansion.

Relative contributions of neutral and niche-based processes to the structure of a desert grassland grasshopper community

Whether neutral or deterministic factors structure biotic communities remains an open question in community ecology. We studied the spatial structure of a desert grassland grasshopper community and tested predictions for species sorting based on niche differentiation (deterministic) and dispersal limitation (neutral). We contrasted the change in species relative abundance and community similarity along an elevation gradient (i.e., environmental gradient) against community change across a relatively homogeneous distance gradient. We found a significant decrease in pairwise community similarity along both elevation and distance gradients, indicating that dispersal limitation plays a role in structuring local grasshopper communities. However, the distance decay of similarity was significantly stronger across the elevational gradient, indicating that niche-based processes are important as well. To further investigate mechanisms underlying niche differentiation, we experimentally quantified the dietary preferences of two common species, Psoloessa texana and Psoloessa delicatula, for the grasses Bouteloua eriopoda and Bouteloua gracilis, which are the dominant plants (~75% of total cover) in our study area. Cover of the preferred host plant explained some of the variation in relative abundances of the two focal species, although much variance in local Psoloessa distribution remained unexplained. Our results, the first to examine these hypotheses in arid ecosystems, indicate that the composition of local communities can be influenced by both probabilistic processes and mechanisms based in the natural histories of organisms.