Constitutive and induced defenses to herbivory in above- and belowground plant tissues

A recent surge in attention devoted to the ecology of soil biota has prompted interest in quantifying similarities and differences between interactions occurring in above- and belowground communities. Furthermore, linkages that interconnect the dynamics of these two spatially distinct ecosystems are increasingly documented. We use a similar approach in the context of understanding plant defenses to herbivory, including how they are allocated between leaves and roots (constitutive defenses), and potential cross-system linkages (induced defenses). To explore these issues we utilized three different empirical approaches. First, we manipulated foliar and root herbivory on tobacco (Nicotiana tabacum) and measured changes in the secondary chemistry of above- and belowground tissues. Second, we reviewed published studies that compared levels of secondary chemistry between leaves and roots to determine how plants distribute putative defense chemicals across the above- and belowground systems. Last, we used meta-analysis to quantify the impact of induced responses across plant tissue types. In the tobacco system, leaf-chewing insects strongly induced higher levels of secondary metabolites in leaves but had no impact on root chemistry. Nematode root herbivores, however, elicited changes in both leaves and roots. Virtually all secondary chemicals measured were elevated in nematode-induced galls, whereas the impact of root herbivory on foliar chemistry was highly variable and depended on where chemicals were produced within the plant. Importantly, nematodes interfered with aboveground metabolites that have biosynthetic sites located in roots (e.g., nicotine) but had the opposite effect (i.e., nematodes elevated foliar expression) on chemicals produced in shoots (e.g., phenolics and terpenoids). Results from our literature review suggest that, overall, constitutive defense levels are extremely similar when comparing leaves with roots, although certain chemical classes (e.g., alkaloids, glucosinolates) are differentially allocated between above- and belowground parts. Based on a meta-analysis of induced defense studies we conclude that: (1) foliar induction generates strong responses in leaves, but much weaker responses in roots, and (2) root induction elicits responses of equal magnitude in both leaves and roots. We discuss the importance of this asymmetry and the paradox of cross-system induction in relation to optimal defense theory and interactions between above- and belowground herbivory.     

Efficacy of stable isotope ratios in assigning endangered migrants to breeding and wintering sites.

A primary constraint on effective conservation of migratory animals is our inability to track individuals through their annual cycle. One such animal is the endangered southwestern subspecies of the Willow Flycatcher, which is difficult to distinguish from conspecifics. Identifying wintering regions used by the endangered subspecies would be an important step in formulating an effective conservation strategy. Our objective was to use stable isotope ratios as a means of identifying wintering sites of Southwestern Willow Flycatchers. We analyzed stable isotope ratios of carbon, nitrogen, and hydrogen from feathers of breeding and wintering Willow Flycatchers. Based on winter samples, we document a positive trend in hydrogen isotope ratios across latitude. We also found that Willow Flycatchers use C4 food webs south of 8 degrees N latitude, but we found no evidence of use of C4 food webs farther north. Nitrogen stable isotope ratios of feathers showed no discernable geographic variation. Discriminant function analyses, based on stable isotope ratios of wintering Willow Flycatchers, were only useful (>50% accurate) for assigning individuals to winter regions if the regions were large and the threshold probability for assignment was relatively high. When using these discriminant functions, most breeding samples of Southwestern Willow Flycatchers were assigned to two wintering regions: central Mexico and Ecuador. We think that assignment of Southwestern Willow Flycatchers to Ecuador is unrealistic. Given the large percentages of samples that could not be classified with certainty, we are not confident that these two regions are truly more likely to harbor wintering Southwestern Willow Flycatchers than other winter regions. We think our inconclusive results are due primarily to weak and nonlinear gradients in isotope ratios across the winter range of Willow Flycatchers.     

Managing plant population spread: prediction and analysis using a simple model

Models can be used to direct the management of population spread for the control of invasives or to encourage species of conservation value. Analytical models are attractive because of their theoretical basis and limited data requirements, but there is concern that their simplicity may limit their practical utility. We address the applied use of simple models in a study of a declining annual herb, Rhinanthus minor. We parameterized a population-spread model using field data on demography and dispersal for four management systems: grazed only (GR), hay-cut once (H1), hay-cut twice (H2), and hay-cut with autumn grazing (HG). Within a replicated experiment we measured spread rates of introduced R. minor populations over eight years. The modeled and measured spread rates were very similar in terms of both patterns of management effects and absolute values, so that in both cases HG > H2, H1 > GR. The treatments affected both dispersal and demography (establishment and survival) and so we used decomposition approaches to analyze the major causes of differences in population spread. Increased dispersal under hay-cutting was more important than demographic changes and accounted for approximately 70% of the differences in spread rate between the hay-cut and grazed-only treatments. Furthermore, management effects on the tail of the dispersal curve were by far the most critical in governing spread. This study suggests that simple models can be used to inform practical conservation management, and we demonstrate straightforward uses of our model to predict the impacts of different management strategies. While simple models can give accurate projections, we emphasize that they must be parameterized with high-quality data gathered at the appropriate spatial scale.     

Forward selection of explanatory variables

This paper proposes a new way of using forward selection of explanatory variables in regression or canonical redundancy analysis. The classical forward selection method presents two problems: a highly inflated Type I error and an overestimation of the amount of explained variance. Correcting these problems will greatly improve the performance of this very useful method in ecological modeling. To prevent the first problem, we propose a two-step procedure. First, a global test using all explanatory variables is carried out. If, and only if, the global test is significant, one can proceed with forward selection. To prevent overestimation of the explained variance, the forward selection has to be carried out with two stopping criteria: (1) the usual alpha significance level and (2) the adjusted coefficient of multiple determination (Ra(2)) calculated using all explanatory variables. When forward selection identifies a variable that brings one or the other criterion over the fixed threshold, that variable is rejected, and the procedure is stopped. This improved method is validated by simulations involving univariate and multivariate response data. An ecological example is presented using data from the Bryce Canyon National Park, Utah, U.S.A.     

Is spread of invasive species regulated? Using ecological theory to interpret statistical analysis

We investigate a recent proposal that invasive species display patterns of spatial "spread regulation" analogous to density-dependent regulation of population abundances. While invasive species do offer valuable tests of ecological theories about spatial spread, we argue that the statistical approach used in the study is not useful, and that the proposed definition of "spread regulation" is likely to be confusing. While concepts of negative feedbacks in spatial spread may be reasonable, the proposed definition of "spread regulation" encompasses accelerating, constant, or decelerating spread. There is no compelling biological or practical reason to adopt such a definition. Moreover, we show that the statistical patterns (from time series of ratios of newly to recently invaded sites) proposed as evidence of spread regulation are predictable from basic diffusion models or other common models of constant spread with some stochasticity in dynamics and/or observations. Because such a wide range of processes would generate the observed patterns, no clear biological conclusions emerge from the proposed approach to spread analysis. When regarded in the context of the impacts and management of invasive species, the proposed regulation concept has the potential to create costly misunderstandings.     

A risk-based screening approach for prioritizing contaminants and exposure pathways at Superfund sites

Contamination at Superfund sites can involve mixtures of chemicals and radionuclides in a variety of environmental media. Determining priorities for evaluation and remediation of various contaminants is an important part of the initial phases of any site investigation. An effective screening analysis at the beginning of the project can help by identifying both those situations in need of immediate remedial attention and those which require further sampling and evaluation. The screening approach discussed here is made up of two sets of calculations designed to provide upper- and lower-bound estimates of health risk to individuals likely to receive the highest exposures. This approach allows rapid identification of contaminants which pose a negligible risk and can be assigned a low priority for remedial attention or which pose a substantial risk and should be given the highest priority for appropriate remediation efforts. Contaminants designated as neither high- nor low-priority should be investigated in more detail prior to making decisions regarding the need for or method of remediation. The utility of this approach has already been demonstrated in the evaluation of contamination in the Clinch and Tennessee River systems originating from historical operations of atomic weapons and energy research facilities near Oak Ridge, Tennessee.     

Importance of eating habits and sample size in the estimation of environmental mercury contamination using biological indicators

The importance of eating habits and the number of fish sampled in the estimation of mercury environmental contamination through biological indicators was studied. The species used were one with piscivorous habits (Anguilla anguilla) and another with omnivorous habits (Ciprinus carpio). From one original sample for each species, randomized samples were generated using the subroutine GGSRS from the mathematics library IMSL, and later the statistical significance of Pearson's correlation coefficient for the relationship between fish weight and mercury concentrations in muscle was obtained for each random sample. The results show that the use of omnivorous species such as Ciprinus carpio as a biological indicator of contamination enables us to carry out estimations with a greater degree of confidence than when piscivorous species such as Anguilla anguilla are used.     

Patchy distributions: Optimising sample size

A method for estimating sample size which does not require an a priori definition of desired precision, or the assumption that the population is normally distributed with constant variance, has recently been proposed. This paper discusses this method and presents five modifications which make the method easier to use and reduce the probability of estimating a larger sample size than is actually required. The method is extended and used to estimate the mean abundance of patchily distributed benthic organisms. The technique can be used to guide the design of any environmental sampling programme, be it physical, chemical or biological, where comparisons between times and/or locations are required. Trade-offs between numbers of replicates and numbers of levels/sites are discussed.     

Effectiveness of Panama as an intercontinental land bridge for large mammals

Habitat fragmentation is a primary driver of wildlife loss, and establishment of biological corridors is a common strategy to mitigate this problem. A flagship example is the Mesoamerican Biological Corridor (MBC), which aims to connect protected forest areas between Mexico and Panama to allow dispersal and gene flow of forest organisms. Because forests across Central America have continued to degrade, the functioning of the MBC has been questioned, but reliable estimates of species occurrence were unavailable. Large mammals are suitable indicators of forest functioning, so we assessed their conservation status across the Isthmus of Panama, the narrowest section of the MBC. We used large-scale camera-trap surveys and hierarchical multispecies occupancy models in a Bayesian framework to estimate the occupancy of 9 medium to large mammals and developed an occupancy-weighted connectivity metric to evaluate species-specific functional connectivity. White-lipped peccary (Tayassu pecari), jaguar (Panthera onca), giant anteater (Myrmecophaga tridactyla), white-tailed deer (Odocoileus virginianus), and tapir (Tapirus bairdii) had low expected occupancy along the MBC in Panama. Puma (Puma concolor), red brocket deer (Mazama temama), ocelot (Leopardus pardalis), and collared peccary (Pecari tajacu), which are more adaptable, had higher occupancy, even in areas with low forest cover near infrastructure. However, the majority of species were subject to ≥1 gap that was larger than their known dispersal distances, suggesting poor connectivity along the MBC in Panama. Based on our results, forests in Darien, Donoso–Santa Fe, and La Amistad International Park are critical for survival of large terrestrial mammals in Panama and 2 areas need restoration.     

A climate-change vulnerability and adaptation assessment for Brazil's protected areas

Brazil hosts the largest expanse of tropical ecosystems within protected areas (PAs), which shelter biodiversity and support traditional human populations. We assessed the vulnerability to climate change of 993 terrestrial and coastal-marine Brazilian PAs by combining indicators of climatic-change hazard with indicators of PA resilience (size, native vegetation cover, and probability of climate-driven vegetation transition). This combination of indicators allows the identification of broad climate-change adaptation pathways. Seventeen PAs (20,611 km²) were highly vulnerable and located mainly in the Atlantic Forest (7 PAs), Cerrado (6), and the Amazon (4). Two hundred fifty-eight PAs (756,569 km²), located primarily in Amazonia, had a medium vulnerability. In the Amazon and western Cerrado, the projected severe climatic change and probability of climate-driven vegetation transition drove vulnerability up, despite the generally good conservation status of PAs. Over 80% of PAs of high or moderate vulnerability are managed by indigenous populations. Hence, besides the potential risks to biodiversity, the traditional knowledge and livelihoods of the people inhabiting these PAs may be threatened. In at least 870 PAs, primarily in the Atlantic Forest and Amazon, adaptation could happen with little or no intervention due to low climate-change hazard, high resilience status, or both. At least 20 PAs in the Atlantic Forest, Cerrado, and Amazonia should be targeted for stronger interventions (e.g., improvement of ecological connectivity), given their low resilience status. Despite being a first attempt to link vulnerability and adaptation in Brazilian PAs, we suggest that some of the PAs identified as highly or moderately vulnerable should be prioritized for testing potential adaptation strategies in the near future.