Heavy browsing affects the hydraulic capacity of Ceanothus rigidus (Rhamnaceae).

Defoliation by herbivores can reduce carbon assimilation, change plant water relations, and even shift the biotic structure of plant communities. In this study, we took advantage of a long-term deer exclosure experiment to examine the consequences of persistent deer herbivory on plant water relations and the xylem structure-function relationships in Ceanothus rigidus, a maritime chaparral shrub in coastal California. Browsed plants had thicker stems with many intertwined short distal twigs, and significantly higher sapwood-to-leaf area ratios than their non-browsed counterparts. Leaf area-specific hydraulic conductivity was similar in both browsed and non-browsed plants, but xylem area-specific conductivity was significantly lower in the browsed plants. Vessel diameters were equivalent in both plant groups, but the number of vessels on a transverse area basis was nearly 40% lower in the browsed plants, accounting for their lower transport efficiency. Mid-day in situ water potentials and losses of hydraulic conductivity due to embolism were similar in both groups of plants but stomatal conductance was higher in the browsed shrubs in the early part of the growing season. We discuss our findings in the context of whole-plant ecophysiology, and explore the consequences of herbivory on hormonal signals, wood anatomy, and xylem function.

Range-expanding populations of a globally introduced weed experience negative plant-soil feedbacks.

BACKGROUND: Biological invasions are fundamentally biogeographic processes that occur over large spatial scales. Interactions with soil microbes can have strong impacts on plant invasions, but how these interactions vary among areas where introduced species are highly invasive vs. naturalized is still unknown. In this study, we examined biogeographic variation in plant-soil microbe interactions of a globally invasive weed, Centaurea solstitialis (yellow starthistle). We addressed the following questions (1) Is Centaurea released from natural enemy pressure from soil microbes in introduced regions? and (2) Is variation in plant-soil feedbacks associated with variation in Centaurea's invasive success? METHODOLOGY/PRINCIPAL FINDINGS: We conducted greenhouse experiments using soils and seeds collected from native Eurasian populations and introduced populations spanning North and South America where Centaurea is highly invasive and noninvasive. Soil microbes had pervasive negative effects in all regions, although the magnitude of their effect varied among regions. These patterns were not unequivocally congruent with the enemy release hypothesis. Surprisingly, we also found that Centaurea generated strong negative feedbacks in regions where it is the most invasive, while it generated neutral plant-soil feedbacks where it is noninvasive. CONCLUSIONS/SIGNIFICANCE: Recent studies have found reduced below-ground enemy attack and more positive plant-soil feedbacks in range-expanding plant populations, but we found increased negative effects of soil microbes in range-expanding Centaurea populations. While such negative feedbacks may limit the long-term persistence of invasive plants, such feedbacks may also contribute to the success of invasions, either by having disproportionately negative impacts on competing species, or by yielding relatively better growth in uncolonized areas that would encourage lateral spread. Enemy release from soil-borne pathogens is not sufficient to explain the success of this weed in such different regions. The biogeographic variation in soil-microbe effects indicates that different mechanisms may operate on this species in different regions, thus establishing geographic mosaics of species interactions that contribute to variation in invasion success.

Growth, fire history, and browsing recorded in wood rings of shrubs in a mild temperate climate.

Separate effects of abiotic and biotic factors on the structure and dynamics of ecological communities may be recorded in growth rings of woody plants. We used Ceanothus cuneatus rigidus and Arctostaphylos pumila to tease apart the roles of fire, rain, and herbivores on the histories and community structure of four areas in a coastal mediterranean-type climate in central California with mild winters and mild summers. Ring widths of both species were related to rainfall in two of the areas; heavy deer browsing on Ceanothus overwhelmed the climate signal in the others. Ceanothus germination was more closely related to heavy rainfall, especially during ENSO years, than to fire events. In a related greenhouse experiment that evaluated these observations, the same proportions of new Ceanothus seeds germinated after burning and after receiving regular water for several months, but germination of old seeds responded primarily to the fire treatment. In areas where heavy browsing by mammals reduces recruitment and growth of Ceanothus and increases mortality, the continuance of the Ceanothus population must rely heavily on germination from the persistent seed bank during unusually wet years or after occasional fires. Because Arctostaphylos can produce new stems from underground roots, individual plants may survive and produce seeds until another fire.

Climatic and biotic stochasticity: disparate causes of convergent demographies in rare, sympatric plants.

Species with known demographies may be used as proxies, or approximate models, to predict vital rates and ecological properties of target species that either have not been studied or are species for which data may be difficult to obtain. These extrapolations assume that model and target species with similar properties respond in the same ways to the same ecological factors, that they have similar population dynamics, and that the similarity of vital rates reflects analogous responses to the same factors. I used two rare, sympatric annual plants (sand gilia [Gilia tenuiflora arenaria] and Monterey spineflower [Chorizanthe pungens pungens]) to test these assumptions experimentally. The vital rates of these species are similar and strongly correlated with rainfall, and I added water and/or prevented herbivore access to experimental plots. Their survival and reproduction were driven by different, largely stochastic factors and processes: sand gilia by herbivory and Monterey spineflower by rainfall. Because the causal agents and processes generating similar demographic patterns were species specific, these results demonstrate, both theoretically and empirically, that it is critical to identify the ecological processes generating observed effects and that experimental manipulations are usually needed to determine causal mechanisms. Without such evidence to identify mechanisms, extrapolations among species may lead to counterproductive management and conservation practices.

Indirect interactions between browsers and seed predators affect the seed bank dynamics of a chaparral shrub.

Interactions between herbivores and seed predators may have long-term consequences for plant populations that rely on persistent seed banks for recovery after unpredictable fires. We assessed the effects of browsing by deer and seed predation by rodents, ants and birds on the densities of seeds entering the seed bank of Ceanothus cuneatus var. rigidus, a maritime chaparral shrub in coastal California. Ceanothus produced many more seeds when protected from browsers in long-term experimental exclosures than did browsed plants, but the seed densities in the soil beneath browsed and unbrowsed Ceanothus were the same at the start of an intensive one-year study. The density of seeds in the soil initially increased in both treatments following summer seed drop: while densities returned to pre-drop levels within a few weeks under browsed plants, soil seed densities remained high for 5-8 months beneath unbrowsed plants. Rodent abundance (especially deer mice) was higher near unbrowsed plants than >30 m away, and rodents removed Ceanothus seeds from dishes in the experimental plots. At least in the short term, rodent density and rates of seed removal were inversely related to the intensity of browsing. Our data have management implications for maintaining viable Ceanothus populations by regulating the intensity of browsing and the timing, intensity and frequency of fires.

Coevolution of Contrary Choices in Host-Parasitoid Systems.

We investigate patch selection strategies of hosts and parasitoids in heterogeneous environments. Previous theoretical work showed that when host traits vary among patches, coevolved populations of hosts and parasitoids make congruent choices (i.e., hosts and parasitoids preferentially select the same patches) and exhibit direct density dependence in the distribution of percent parasitism. However, host-parasitoid systems in the field show a range of patterns in percent parasitism, while behavioral studies indicate that hosts and parasitoids can exhibit contrary choices (i.e., hosts avoid patches favored by the parasitoid). We extend previous theory by permitting life-history traits of the parasitoid as well as the host to vary among patches. Our analysis implies that in coevolutionarily stable populations, hosts preferentially select patches that intrinsically support higher host equilibrium numbers (i.e., the equilibrium number achieved by hosts when both populations are confined to a single patch) and that parasitoids preferentially select patches that intrinsically support higher parasitoid equilibrium numbers (i.e., the equilibrium number achieved by the parasitoids when both populations are confined to a patch). Using this result, we show how variation in life-history traits among patches leads to contrary or congruent choices or leads to direct density dependence, inverse density dependence, or density independence in the distribution of percent parasitism. In addition, we determine when populations playing the coevolutionarily stable strategies are ecologically stable. Our analysis shows that heterogeneous environments containing patches where the intrinsic rate of growth of the host and the survivorship rate of the parasitoid are low result in the coevolved populations exhibiting contrary choices and, as a result, promote ecological stability.

Direct and indirect effects of climate change on St John's wort, Hypericum perforatum L. (Hypericaceae).

We report results from a continuing, long-term field experiment addressing biotic responses to climatic change in grasslands. We focus on effects of summer precipitation (enhanced rainfall, drought, control) and winter ground temperatures (warming, control) on growth, reproduction and herbivory in St John's wort, Hypericum perforatum L. Both winter warming and summer rainfall regimes modified performance and interactions of H. perforatum, particularly those with herbivorous insects. Winter warming had positive effects, with earlier initiation of plant growth and reduced damage by gall-forming and sucking insects in spring, but also had strong negative effects on plant height, flowering, and reproduction. Summer drought reduced reproductive success, but even severe drought did not affect plant growth or flowering success directly. Rather, summer drought acted indirectly by modifying interactions with herbivorous insects via increased vulnerability of the plants to herbivory on flowers and capsules. Overall, the effects of summer precipitation were expressed mainly through interactions that altered the responses to increased winter temperatures, particularly as summer drought increased. The field site, in Oxfordshire, UK, is near the northern limit of distribution of the species, and the experiment tested probable responses of H. perforatum as climates shift towards those more typical of the current center of the distribution of the species. However, if climates do change according to the projected scenarios, then H. perforatum is unlikely to fare well near its northern boundary. Increased winter temperatures, particularly if accompanied by increased summer drought, will probably render this species even less abundant in England than at present.

Effects of mammal and insect herbivory on population dynamics of a native Californian thistle, Cirsium occidentale.

We explored consequences of spatial and temporal heterogeneity in herbivory on the survival, growth, and reproduction of the Californian native dune thistle, Cirsium occidentale, in coastal and inland sites, for 2 years. We assessed the relative impacts of insect and mammalian herbivores and compared the relative importance of herbivory in coastal and inland habitats and among locations with different microclimates across a coastal dune. Effects of insect and mammalian herbivores were tested with a combination of insecticidal spray and cage exclusion treatments in a factorial experiment at the coastal site. Mammalian herbivores strongly affected the population dynamics of C. occidentale in both years, and their effects were augmented by fungal infection (1991), herbivory by stem-borers (1990) and, to a lesser extent, by insect seed predators in both years. Mammals caused most plant deaths, but the mammal species responsible differed among sites. Rabbit herbivory altered the vegetative growth of coastal thistles and significantly modified other key aspects of Cirsium demography, including growth rate and timing of reproduction. Small, uncaged plants grazed by rabbits took at least 1 year longer to mature than did caged plants. Larvae of Pyrausta subsequalis were the only insects that killed established plants. In 1990 and 1991, the numbers of insects damaging seed heads before dispersal were low, but were sufficient to cause receptacle and seed damage. The number of mature, undamaged seeds (and percent successful seed production) was reduced significantly only for heads infested by fungi near the ocean in 1991: the fungus occurred in 37% of heads and caused a 77% reduction in mature seeds.

Eucalypt responses to fertilization and reduced herbivory.

We manipulated soil fertility and insect attack for two species of Eucalyptus in natural stands of subalpine woodland on shallow, infertile granitic soils. E. pauciflora and E. stellulata responded in similar ways to simultaneous insecticide and fertilizer treatments. Eliminating herbivorous insects produced the largest changes - improved plant growth, increased leaf N and P, and reduced leaf specific density. Fertilizer regime modified some leaf properties, but had little effect on tree growth. E. stellulata trees were initially shorter than E. pauciflora, but grew faster without herbivores; by the end of the experiment both species were the same size when herbivores were removed. Foliage N and P levels increased most in trees with the most balanced fertilizer addition (NPK), and increased in all trees protected from insects, regardless of fertilizer regime. In this system, herbivorous insects exacerbated the effects of nutrientpoor soils, and may affect dominance of Eucalyptus species in mature forests.

Parasitism rates and sex ratios of a parasitoid wasp: effects of herbivore and plant quality.

We studied interactions among collards, Brassica oleracea var. acephala, the diamondback moth (DBM), Plutella xylostella (Lepidoptera: Yponomeutidae) and its parasitoid Diadegma insulare (Hymenoptera: Ichneumonidae) by manipulating plant nitrogen (N) concentrations in field and laboratory experiments. Parasitoid abundance strongly reflected DBM abundance and was related to total leaf N. Parasitism rates were high (70.7%) and density-independent. Wasp sex ratios varied markedly (3-93% female) in response to the herbivores, the plants, or both. Higher proportions of female wasps emerged from DBM larvae on plants with high leaf N than on unfertilized plants. More female wasps also emerged from larvae parasitized as larger instars. We suggest that wasps have the potential to control DBM populations through long-term numerical responses mediated by variable sex ratios.

Effects of variation in Eucalyptus essential oil yield on insect growth and grazing damage.

Levels of insect attack and yields of leaf essential oils in Eucalyptus vary widely within and among species. We tested the hypotheses that 1) metabolic cost of oil detoxification increases with increasing oil yield, resulting in lower herbivore growth rates and, consequently, 2) in lower herbivore damage to plants. Distribution of insect damage, eggs, immature insects and adults and feeding rates, growth and survivorship of insects do not support the hypotheses, although a threshold level of oil may be necessary to influence herbivorous insects. Herbivorous beetles tested do not detoxify essential oils. Levels of leaf nitrogen, rather than oil content, explained differences in insect feeding and growth.

Insect grazing on Eucalyptus in response to variation in leaf tannins and nitrogen.

Many species of Eucalyptus, one of the dominant genera in Australian forests and woodlands, contain high levels of tannins and other phenols and are also heavily damaged by grazing insects. These phenols do not appear to affect insect attack because a wide range of concentrations of condensed tannins and other phenols in leaves of 13 Eucalyptus sp. influenced neither feeding rates of Paropsis atomaria larvae, nor their nitrogen use efficiencies. We discuss reasons why tannins may not appreciably reduce the availability of nitrogen (N) to these insects. Performance was directly related to leaf N concentration, and growth rates, N gains, and N use efficiencies all increased as leaf N content increased, although absolute feeding rates remained constant. These relationships differ from those found in insects feeding on other plants, and we suggest that the low N contents common in Eucalyptus leaves may be responsble. We propose that the extensive damage observed in many eucalypts is in part related to the high feeding rates maintained by individual larvae.