Latitudinal variation in top-down and bottom-up control of a salt marsh food web.

The shrub Iva frutescens, which occupies the terrestrial border of U.S. Atlantic Coast salt marshes, supports a food web that varies strongly across latitude. We tested whether latitudinal variation in plant quality (higher at high latitudes), consumption by omnivores (a crab, present only at low latitudes), consumption by mesopredators (ladybugs, present at all latitudes), or the life history stage of an herbivorous beetle could explain continental-scale field patterns of herbivore density. In a mesocosm experiment, crabs exerted strong top-down control on herbivorous beetles, ladybugs exerted strong top-down control on aphids, and both predators benefited plants through trophic cascades. Latitude of plant origin had no effect on consumers. Herbivorous beetle density was greater if mesocosms were stocked with beetle adults rather than larvae, and aphid densities were reduced in the "adult beetle" treatment. Treatment combinations representing high and low latitudes produced patterns of herbivore density similar to those in the field. We conclude that latitudinal variation in plant quality is less important than latitudinal variation in top consumers and competition in mediating food web structure. Climate may also play a strong role in structuring high-latitude salt marshes by limiting the number of herbivore generations per growing season and causing high overwintering mortality.

Variation in tolerance and resistance to the leafhopper Empoasca fabae (Hemiptera: Cicadellidae) among potato cultivars: implications for action thresholds.

The potato leafhopper, Empoasca fabae (Harris) (Hemiptera: Cicadellidae), is an emerging pest of potato and insecticide applications to control this insect have increased in recent years. Based on field observations of leafhopper-crop dynamics, however, currently recommended action thresholds seem to be overly conservative. As a result, we initiated two experiments designed to quantify the impact of leafhoppers on potato yield, and determine how the magnitude of this effect changes among cultivars. In experiment 1, leafhoppers were manipulated (control versus insecticide-treated plots) on 17 potato varieties. In experiment 2, three cultivars (Superior, Atlantic, and Snowden) were planted representing early-, mid-, and late-season maturing lines, and six insecticide spray regimes were imposed (early-, late-, and full-season applications at high and low rates). In both experiments, leafhopper abundance, plant damage, and potato yield were measured. Overall, leafhoppers reduced yield in control plots by 15.7% relative to insecticide-treated plots. Leafhopper impact, however, varied among cultivars; a significant effect of leafhoppers on yield was detected in 6, 12, and 59% of cultivars tested in each of three trials. Of the 44 cases in which leafhoppers exceeded action thresholds, yield loss was only documented in 13 cases. Data from these experiments provide evidence that such variable effects ofleafhoppers on yield are explained by cultivar-specific resistance and tolerance traits. Our results suggest that potato growers can accept higher leafhopper densities than current thresholds recommend, particularly when cultivating resistant and/or tolerant varieties.

Nutritional constraints in terrestrial and freshwater food webs.

Biological and environmental contrasts between aquatic and terrestrial systems have hindered analyses of community and ecosystem structure across Earth's diverse habitats. Ecological stoichiometry provides an integrative approach for such analyses, as all organisms are composed of the same major elements (C, N, P) whose balance affects production, nutrient cycling, and food-web dynamics. Here we show both similarities and differences in the C:N:P ratios of primary producers (autotrophs) and invertebrate primary consumers (herbivores) across habitats. Terrestrial food webs are built on an extremely nutrient-poor autotroph base with C:P and C:N ratios higher than in lake particulate matter, although the N:P ratios are nearly identical. Terrestrial herbivores (insects) and their freshwater counterparts (zooplankton) are nutrient-rich and indistinguishable in C:N:P stoichiometry. In both lakes and terrestrial systems, herbivores should have low growth efficiencies (10-30%) when consuming autotrophs with typical carbon-to-nutrient ratios. These stoichiometric constraints on herbivore growth appear to be qualitatively similar and widespread in both environments.

The influence of dispersal and diet breadth on patterns of genetic isolation by distance in phytophagous insects.

To determine the effects of dispersal ability and diet breadth on population-genetic structure, we reviewed the allozyme literature and estimated genetic isolation by distance (IBD) for 43 species/host races of phytophagous insects. Subsequently, we tested two opposing hypotheses regarding the influence of dispersal ability on IBD: that IBD slopes do not vary with mobility, but that intercepts increase with mobility, and, alternatively, that IBD slopes vary with dispersal ability. We found that from tens of kilometers to more than 1,000 km, IBD is weak in sedentary and highly mobile species but pronounced in moderately mobile species. We attribute the weak IBD in strong dispersers to the homogenizing effects of gene flow, whereas in sedentary species, limited gene flow allows nearly all populations to diverge. In intermediate dispersers, genetic homogeneity is achieved at small spatial scales, but limited dispersal promotes genetic divergence over long distances. We also tested the hypothesis that IBD increases with decreasing diet breadth. We discovered no such pattern, casting doubt on the supposition that specialization promotes speciation by influencing population-genetic subdivision. Finally, we found that the number of populations is a more important consideration than the number of polymorphic loci in studies of IBD.

Physiology and ecology of dispersal polymorphism in insects.

Studies of dispersal polymorphism in insects have played a pivotal role in advancing our understanding of population dynamics, life history evolution, and the physiological basis of adaptation. Comparative data on wing-dimorphic insects provide the most definitive evidence to date that habitat persistence selects for reduced dispersal capability. The increased fecundity of flightless females documents that a fitness trade-off exists between flight capability and reproduction. However, only recently have studies of nutrient consumption and allocation provided unequivocal evidence that this fitness trade-off results from a trade-off of internal resources. Recent studies involving wing-dimorphic insects document that flight capability imposes reproductive penalties in males as well as females. Direct information on hormone titers and their regulation implicates juvenile hormone and ecdysone in the control of wing-morph determination. However, detailed information is available for only one species, and the physiological regulation of wing-morph production remains poorly understood. Establishing a link between the ecological factors that influence dispersal and the proximate physiological mechanisms regulating dispersal ability in the same taxon remains as a key challenge for future research.

Differences in environmental response between the sexes of the dioecious shrub Baccharis halimifolia (Compositae).

Baccharis halimifolia (Compositae) is a dioecious shrub which grows on the upland fringe of tidal marshes along the Atlantic and Gulf Coasts of North America. We examined the responses of the two sexes to variation in nutrient and moisture availability plant density, and defoliation. By growing plants from seedlings to flowering adults under various combinations of soil type, fertilization rate and plant density, we were able to establish different rates of plant growth and mortality. Plants grown at high density and low nutrient and water supply grew the least, incurrent the most mortality and showed a male-biased sex ratio (73% male). At low density with abundant nutrients and water, plants grew more, survived well, flowered frequently, and were female-biased (75% female). Changes in sex ratio were probably the result of sex-related mortality rather than sexual lability of the seedlings. While changes in sex ratio occurred under experimental conditions in the green-house, no evidence for differences in habitat utilization between the sexes were found in the field and the sex ratio (59% female) did not vary across habitats. In the marsh habitats we sampled where water and nutrients were apparently available, there was no evidence for differential mortality between the sexes. When defoliated (75% of leaf tissue), both sexes showed similar reductions in reproductive effort (number of flower heads/shoot). Our results indicate that differences between the sexes of Baccharis in their response to environmental growing conditions is an important ecological factor associated with the separation of male and female function into separate individuals.

Patterns of growth, reproduction, defense, and herbivory in the dioecious shrub Baccharis halimifolia (Compositae).

Patterns of growth, reproduction, defense (leaf resin) and herbivory were compared between the sexes of the dioecious shrub Baccharis halimifolia (Compositae). Male plants possessed longer shoots and more tender leaves, grew faster, and flowered and senesced earlier than female plants. Levels of leaf nitrogen, water content, and acetone-soluble resin (shown to deter feeding by polyphagous insect herbivores) did not differ between male and female plants. When offered a choice between leaves from male and female plants, adults of two leaf beetles (Chrysomelidae), the monophagous Trirhabda bacharidis and the polyphagous Paria thoracica, both preferred to feed on male leaves. Similarly, the daily fecundity of older females of T. bacharidis was higher when they were fed leaves from male compared to female plants. However, adult survivorship and total fecundity of T. baccharidis did not differ between male and female leaf treatments. We attribute the feeding preference for and slight increase in fecundity on male plants to the tenderness of male leaves. Larvae of the fly Tephritis subpura (Tephritidae) fed exclusively in the sterile receptacle of male flower heads (85% infested), but the phenology was such that pollen production was not adversely affected. Larvae of two other flies Dasineura sp. and Contarinia sp. (Cecidomyiidae) occupied >95% of only female flower heads where they fed among and on the developing seeds. We conclude that foliage-feeding herbivores are unlikely candidates to explain the female-biased sex ratio (59% female) of B. halimifolia plants in the field, and that their preference for male plants is a result of plant characteristics (e.g. rapid growth) that have been selected by some other factor. However, our data on selective floral herbivory in B. halimifolia are in accord with the argument that dioecy reduces the inadvertent loss of flower parts of one sex when herbivores feed on flower parts of the opposite sex.