Multiseasonal management of an agricultural pest. I: Development of the theory

A framework for analyzing the trade-off between economic yield from a crop and buildup of resistance to pesticide caused by repeated applications of pesticide is developed. The analysis begins with the case of age-independent pest dynamics, in which pests infest a field by arriving from an external pool. Initially, it is assumed that the pest genetics of interest are single locus, two allele, with resistance to pesticide dominant and susceptible pests more fit in the absence of spraying. The pesticide is applied only once during the season, with timing and intensity of the application as control variables. Interseasonal pest and crop dynamics are studied by solving appropriate ordinary differential equations. Intraseasonal pest dynamics are assumed to follow the Hardy-Weinberg formula. It is shown that the three class diploid model can be replaced by a two class haploid model with essentially no change in the results. A model based on partial differential equations is developed, for the case in which pest dynamics depend upon age, and it is shown that the partial differential equation model can be replaced by a pair of coupled ordinary differential equations. The main operational conclusion in this paper is that the timing of the application of pesticide can be used to control buildup of resistance and that the intensity of the application can be used to control the crop yield.     

On the optimal allocation of pesticides with increasing resistance: The case of alfalfa weevil

The problem of pesticide application under increasing pesticide resistance is explored. A theoretical model is developed to determine optimal pesticide use. This allocation is compared to the laissez-faire solution and to the centralized solution with incomplete information about pesticide resistance. The methodology is then applied in a case study on the Egyptian alfalfa weevil in California.     

Mitigation of agricultural nonpoint-source pesticide pollution in artificial wetland ecosystems

Contamination caused by pesticides in agriculture is a source of environmental poor water quality in some of the European Union countries. Without treatment or targeted mitigation, this pollution is diffused in the environment. Pesticides and some metabolites are of increasing concern because of their potential impacts on the environment, wildlife and human health. Within the context of the European Union (EU) water framework directive context to promote low pesticide-input farming and best management practices, the EU LIFE project ArtWET assessed the efficiency of ecological bioengineering methods using different artificial wetland (AW) prototypes throughout Europe. We optimized physical and biological processes to mitigate agricultural nonpoint-source pesticide pollution in artificial wetland ecosystems. Mitigation solutions were implemented at full-scale demonstration and experimental sites. We tested various bioremediation methods at seven experimental sites. These sites involved (1) experimental prototypes, such as vegetated ditches, a forest microcosm and 12 wetland mesocosms, and (2) demonstration prototypes: vegetated ditches, three detention ponds enhanced with technology of constructed wetlands, an outdoor bioreactor and a biomassbed. This set up provides a variety of hydrologic conditions, with some systems permanently flooded and others temporarily flooded. It also allowed to study the processes both in field and controlled conditions. In order to compare the efficiency of the wetlands, mass balances at the inlet and outlet of the artificial wetland will be used, taking into account the partition of the studied compound in water, sediments, plants, and suspended solids. The literature background necessary to harmonize the interdisciplinary work is reviewed here and the theoretical framework regarding pesticide removal mechanisms in artificial wetland is discussed. The development and the implementation of innovative approaches concerning various water quality sampling strategies for pesticide load estimates during flood, specific biological endpoints, innovative bioprocess applied to herbicide and copper mitigation to enhance the pesticide retention time within the artificial wetland, fate and transport using a 2D mixed hybrid finite element model are introduced. These future results will be useful to optimize hydraulic functioning, e.g., pesticide resident time, and biogeochemical conditions, e.g., dissipation, inside the artificial wetlands. Hydraulic retention times are generally too low to allow an optimized adsorption on sediment and organic materials accumulated in artificial wetlands. Absorption by plants is not either effective. The control of the hydraulic design and the use of adsorbing materials can be useful to increase the pesticides residence time and the contact between pesticides and biocatalyzers. Pesticide fluxes can be reduced by 50–80% when hydraulic pathways in artificial wetlands are optimized by increasing ten times the retention time, by recirculation of water, and by deceleration of the flow. Thus, using a bioremediation method should lead to an almost complete disappearance of pesticides pollution. To retain and treat the agricultural nonpoint-source po a major stake for a sustainable development.     

Sublethal pesticide concentrations and predation jointly shape life history: behavioral and physiological mechanisms.

Despite their relevance for risk assessment, the interactive effects of pesticide and predation cues are poorly understood because the underlying behavioral and physiological mechanisms are largely unknown. To explore these mechanisms, we reared larvae of the damselfly Coenagrion puella at three different predation risk levels and a range of environmentally realistic concentrations of three pesticides used worldwide (atrazine, carbaryl, and endosulfan). We compared key development responses (growth rate, developmental time, and final size) against food ingestion, assimilation, and conversion efficiency, and acetylcholinesterase (AChE) activity. Predation risk impaired all endpoints, including AChE activity, while the effects of pesticide stress were smaller for atrazine and endosulfan and absent for carbaryl. The effects of both stressors and their interaction on life history were mostly indirect through resource acquisition and energy allocation. Compensatory physiological mechanisms to pesticide stress (atrazine and endosulfan) were present in larvae reared in the absence of predation stress but were offset under predation stress. As a result, smaller size (atrazine and endosulfan) and lower growth rate (endosulfan) from pesticide stress were only found in the highest predation risk treatment. Our results provide insight as to the conditions under which interactions between stressors are likely to occur: damselfly populations at high density and living in fish ponds will be more affected by pesticides than populations at low densities in fishless ponds. By identifying variables that may shape the interaction between predation stress and other stressors such as pesticides, our mechanistic approach may help to bridge the gap between laboratory and field studies.     

涕灭威农药污染地下水的影响因子分析

农药在土壤中的淋溶受许多因子的影响与制约。本文根据农药在土壤中的环境行为，建立了农药土壤残留动态与淋溶归宿的计算机模型，并用此模型计算了模拟各种不同环境条件下涕灭威农药在土层中的淋溶情况。结果表明：农药的使用量、土层中农药的降解半衰期、施药地区土壤的质地、降水或灌溉水量及其距施药后的时间对涕灭威在土层中的淋溶有较大的影响，而土壤有机碳含量则影响不大；地下水埋深对农药地下水污染的影响颇大。地下水埋深不足１ｍ的地区，极易受农药的污染。     

Modeling pesticide fate in a small tidal estuary

The exposure analysis modeling system (EXAMS), a pesticide fate model developed by the U.S. Environmental Protection Agency, was modified to model the fate of the herbicides atrazine and metolachlor in a small tidally dominated estuary (Bath Creek) in North Carolina, USA where freshwater inflow accounts for only 3% of the total flow. The modifications simulated the changes that occur during the tidal cycle in the estuary, scenarios that are not possible with the original EXAMS model. Two models were created within EXAMS, a steady-state model and a time-variant tidally driven model. The steady-state model accounted for tidal flushing by simply altering freshwater input to yield an estuary residence time equal to that measured in Bath Creek. The tidal EXAMS model explicitly incorporated tidal flushing by modifying the EXAMS code to allow for temporal changes in estuary physical attributes (e.g., volume). The models were validated with empirical measurements of atrazine and metolachlor concentrations in the estuary shortly after herbicide application in nearby fields and immediately following a rain event. Both models provided excellent agreement with measured concentrations. The steady-state EXAMS model accurately predicted atrazine concentrations in the middle of the estuary over the first 3 days and under-predicted metolachlor by a factor of 2–3. The time-variant, tidally driven EXAMS model accurately predicted the rise and plateau of both herbicides over the 6-day measurement period. We have demonstrated the ability of these modified EXAMS models to be useful in predicting pesticide fate and exposure in small tidal estuaries. This is a significant improvement and expansion of the application of EXAMS, and given the wide use of EXAMS for surface water quality modeling by both researchers and regulators and the ability of EXAMS to interface with terrestrial models (e.g., pesticide root zone model) and bioaccumulation models, we now have an easily-accessible and widely accepted means of modeling chemical fate in estuaries.     

Market design in cap and trade programs: Permit validity and compliance timing

Cap and trade programs have considerable heterogeneity in permit validity and compliance timing. For example, permits have different validities across time (e.g., banking, borrowing, and seasons) and space (e.g., zonal restrictions), and compliance timing can be annual, in overlapping cycles, or in multi-year periods. We discuss nine prominent cap and trade programs along these dimensions and construct a general model of permit validity and compliance timing. We derive sufficient conditions under which abatement is invariant to compliance timing, i.e., compliance timing cannot smooth abatement cost shocks. Under these conditions, (i) expected compliance costs are invariant, (ii) the variance of compliance costs increases with the delayed compliance, (iii) equilibrium prices may not be unique, and (iv) the delayed compliance equilibrium may rely on “degenerate” prices not determined by marginal abatement costs. We demonstrate the model's broad applicability by illustrating different types of temporal permit validity.     

Host plant resistance to insects: an eco-friendly approach for pest management and environment conservation

Host plant resistance (HPR) to insects is an effective, economical, and environment friendly method of pest control. The most attractive feature of HPR is that farmers virtually do not need any skill in application techniques, and there is no cash investment by the resource poor farmers. Considerable progress has been made in identification and development of crop cultivars with resistance to the major pests in different crops. There is a need to transfer resistance genes into high-yielding cultivars with adaptation to different agro-ecosystems. Resistance to insects should form one of the criteria to release varieties and hybrids for cultivation by the farmers. Genes from the wild relatives of crops, and novel genes, such as those from Bacillus thuringiensis can also be deployed in different crops to make HPR an effective weapon to minimize the losses due to insect pests. HPR will not only cause a major reduction in pesticide use and slowdown the rate of development of resistance to insecticides in insect populations, but also lead to increased activity of beneficial organisms and reduction in pesticide residues in food and food products.     

Long-term dynamics of biotic and abiotic resistance to exotic species invasion in restored vernal pool plant communities

Invasion of native ecosystems by exotic species can seriously threaten native biodiversity, alter ecosystem function, and inhibit conservation. Moreover, restoration of native plant communities is often impeded by competition from exotic species. Exotic species invasion may be limited by unfavorable abiotic conditions and by competition with native species, but the relative importance of biotic and abiotic factors remains controversial and may vary during the invasion process. We used a long-term experiment involving restored vernal pool plant communities to characterize the temporal dynamics of exotic species invasion, and to evaluate the relative support for biotic and abiotic factors affecting invasion resistance. Experimental pools (n=256) were divided among controls and several seeding treatments. In most treatments, native vernal pool species were initially more abundant than exotic species, and pools that initially received more native seeds exhibited lower frequencies of exotic species over time. However, even densely seeded pools were eventually dominated by exotic species, following extreme climatic events that reduced both native and exotic plant densities across the study site. By the sixth year of the experiment, most pools supported more exotics than native vernal pool species, regardless of seeding treatment or pool depth. Although deeper pools were less invaded by exotic species, two exotics (Hordeum marinum and Lolium multiflorum) were able to colonize deeper pools as soon as the cover of native species was reduced by climatic extremes. Based on an information-theoretic analysis, the best model of invasion resistance included a nonlinear effect of seeding treatment and both linear and nonlinear effects of pool depth. Pool depth received more support as a predictor of invasion resistance, but seeding intensity was also strongly supported in multivariate models of invasion, and was the best predictor of resistance to invasion by H. marinum and L. multilorum. We conclude that extreme climatic events can facilitate exotic species invasions by both reducing abiotic constraints and weakening biotic resistance to invasion.     

A simulation model of population dynamics of the codling moth, Cydia pomonella

A simulation model has been developed that predicts numbers and phenology of a population of codling moth, Cydia pomonella (L.), in an apple orchard. The model is a general insect population model based on the interative-cohort technique. It operates at two time scales: a fine time scale (1 h) for temperature-dependent physiological processes, and a coarse time scale (1 day) for population processes. The population is divided into a specifiable number of stages, and each stage is described by four process functions, which may be of any convenient mathematical form, and may differ among stages. Each stage is divided into cohorts of individuals born or emerged on the same day, and individuals within a cohort are considered probabilistically identical. The model simulates the processes of development, transition among stages, and mortality by using probability distributions representing these processes, and incorporates the effects of pesticides on mortality of the insect. Model output was evaluated by comparison with records of pheromone trap catches of codling moths in commercial apple orchards in North Carolina. The model predicts timing of the first spring flight well, depending on the initial age distribution used. Time between peaks of numbers of adults in the model is about 15 days longer than the observed period between flight peaks in orchards. Sensitivity analysis indicates that this discrepancy may be related to differences between measured ambient temperature and tree canopy temperature. The sensitivities of numbers of insects produced by the model, and timing of peaks in numbers present were determined for each of the parameters in the model. The parameters with greatest effect on the model output were those which control the locations of developmental rate functions and survival functions on the temperature scale. In the model, pesticides had a much larger effect on numbers of adults present than records of moths caught in pheromone traps indicate actually occurred, suggesting that moths caught in traps in commercial orchards where effective pesticides are applied may be largely immigrants.     

杀虫剂敌敌畏对黑眶蟾蜍蝌蚪的毒性影响

采用静态换水法研究了杀虫剂敌敌畏（dichlorvos，DDVP）对黑眶蟾蜍（Bufo melanostictus）蝌蚪的急性毒性和生长发育、生理活动的影响．结果表明，DDVP对黑眶蟾蜍蝌蚪的96h半数致死浓度（LC50，96h）为51．61mg／L．暴露于2．24mg／L的DDVP溶液中10d和20d，蝌蚪的体长日增长率分别降低24．9％和9．5％，并引起黑眶蟾蜍蝌蚪酯酶同工酶（EST）酶带数和强弱发生改变，抑制了蝌蚪的酯酶同工酶活性，而超氧化物歧化酶（SOD）比活受到的影响相对较小．暴露于6．72mg／L的DDVP溶液中10d和20d，蝌蚪的体长日增长率分别降低27．0％和38．8％，酯酶同工酶（EST）酶带的强弱发生明显的改变，SOD酶比活明显低于对照组．说明杀虫剂敌敌畏对黑眶蟾蜍蝌蚪可造成一定的毒性影响．图2表4参22     

土壤中结合残留态农药的生态环境效应

土壤中农药结合残留态的形成，导致其活性暂时降低，但并未从土壤中消失，在特定的环境条件下又重新释放到环境中并表现出较高的生物有效性，从而威胁农产品质量安全与环境质量。文章论述了土壤中结合残留态农药的定义、形成过程及影响因素、老化和释放过程及机制。土壤中结合残留态农药主要通过吸附过程、化学反应及物理镶嵌等作用而形成，其形成过程受农药的结构和化学特性、土壤理化性质、环境条件和农艺措施的影响。老化是化合物和土壤组分紧密结合，减少被普通提取方法提取出来的数量，降低了化合物的生物有效性。同时老化的物质在土壤环境条件改变的情况下又重新释放到土壤溶液中或进行矿化，此过程可以通过物理一化学机制或生物化学作用而发生。重新释放到环境中的结合残留态农药又表现出较高的生物有效性，可能被植物、动物或微生物所吸收，并沿食物链富集和放大或进入水体污染水产品和造成水质恶化，从而威胁人体健康。文章还分析了土壤结合残留态农药可能带来的环境问题，提出了此问题今后研究的方向。     

Pesticide prediction in ground water in North Carolina domestic wells using artificial neural networks

This paper illustrates the application of artificial neural networks (ANN) for prediction of pesticide occurrence in rural domestic wells from the available limited information. Among the three ANN models (a feed-forward back propagation [BP], a radial basis function [RBF] and an adaptive neural network-based fuzzy inference system [ANFIS]) employed for this investigation, the BP neural network was found to be superior to RBF and ANFIS type networks for the detection of pesticide occurrences in wells. For improved model prediction efficiency, optimization of network structure (e.g., number of hidden layers and number of nodes in each hidden layer) and spread (the width of the radial basis function) are important for BP and RBF type of network, respectively. A four layer BP network with a 3:2 neurons ratio of the first hidden layer to the second hidden layer produced better prediction performance efficiencies in terms of the pesticide detection efficiency (E_f), the root mean square error (RMSE), and the correlation coefficient (R) and the overall E_f of the BP neural network was found greater than 85%. Sensitivity analysis was performed to measure the relative importance of one input parameter over the other in pesticide occurrence in wells. It was shown in terms of the prediction efficiencies (E_f, RMSE, and R) of a four-layer BP neural network that the time of sample collection (TSC; month of the year), the depths of wells, and pesticide travel times (PTT) were more important parameters in the prediction of the pesticide occurrences in rural domestic wells. This means that the wells having shallow ground water table are more vulnerable to pesticide occurrence.     

Testing the evolution of increased competitive ability (EICA) hypothesis in a novel framework

The "evolution of increased competitive ability" (EICA) hypothesis proposes that escape from natural enemies, e.g., after transcontinental introductions, alters the selection regime because costly defenses no longer enhance fitness. Such an evolutionary loss of defenses enables resources to be directed toward growth or other traits improving performance. We tested the EICA hypothesis in a novel framework in which the natural enemy is the traveler that follows its widespread host by accidental or deliberate (biocontrol) introductions. In a greenhouse experiment we used populations of Senecio vulgaris from North America, Europe, and Australia that differ in the history of exposure to the rust fungus Puccinia lagenophorae. Contrary to what is predicted by EICA, we found no evidence for increased levels of resistance to the rust fungus in plant populations with a longer history of rust exposure (Australia). Similarly, there was no evidence for reduced fecundity in these populations, although vegetative vigor, measured as secondary branching and growth rate, was lower. The maintenance of high rust resistance in populations with no (North America) or only a short history (Europe) of rust exposure is surprising given that resistance seems to incur considerable fitness costs, as indicated by the negative association between family mean resistance and fitness in the absence of disease observed for all three continents. The comparison of population differentiation in quantitative traits with estimates of differentiation in amplified fragment length polymorphic (AFLP) markers suggests that a number of fitness-related traits are under divergent selection among the studied populations. The proposed framework to test changes in the evolutionary trajectory underlying EICA can be employed in an expanded range of systems. These may include investigations on a cosmopolitan weed or crop when an antagonist is expanding its geographic range (such as our study), studies along a chronosequence of introduction time with expected increasing accumulation of natural enemies over time, or comparisons between introduced plant populations that differ in exposure time to biocontrol organisms.     

Pesticide resistance externalities and optimal mosquito management

A simultaneous equation model of the behavior of a mosquito abatement district based on biological and economic data is presented. Results indicate high long term costs if heavy reliance on chemical pesticide control methods continue, due to a pesticide resistance buildup in the mosquito populations. Physical source reduction methods were shown to be more efficient both in the short and long run. A linear programming model is presented which optimizes the inix of chemical and physical control methods. Results indicate increasing costs of mosquito abatement as pesticide effectiveness declines.     

Abiotic constraints eclipse biotic resistance in determining invasibility along experimental vernal pool gradients.

Effective management of invasive species requires that we understand the mechanisms determining community invasibility. Successful invaders must tolerate abiotic conditions and overcome resistance from native species in invaded habitats. Biotic resistance to invasions may reflect the diversity, abundance, or identity of species in a community. Few studies, however, have examined the relative importance of abiotic and biotic factors determining community invasibility. In a greenhouse experiment, we simulated the abiotic and biotic gradients typically found in vernal pools to better understand their impacts on invasibility. Specifically, we invaded plant communities differing in richness, identity, and abundance of native plants (the "plant neighborhood") and depth of inundation to measure their effects on growth, reproduction, and survival of five exotic plant species. Inundation reduced growth, reproduction, and survival of the five exotic species more than did plant neighborhood. Inundation reduced survival of three species and growth and reproduction of all five species. Neighboring plants reduced growth and reproduction of three species but generally did not affect survival. Brassica rapa, Centaurea solstitialis, and Vicia villosa all suffered high mortality due to inundation but were generally unaffected by neighboring plants. In contrast, Hordeum marinum and Lolium multiflorum, whose survival was unaffected by inundation, were more impacted by neighboring plants. However, the four measures describing plant neighborhood differed in their effects. Neighbor abundance impacted growth and reproduction more than did neighbor richness or identity, with growth and reproduction generally decreasing with increasing density and mass of neighbors. Collectively, these results suggest that abiotic constraints play the dominant role in determining invasibility along vernal pool and similar gradients. By reducing survival, abiotic constraints allow only species with the appropriate morphological and physiological traits to invade. In contrast, biotic resistance reduces invasibility only in more benign environments and is best predicted by the abundance, rather than diversity, of neighbors. These results suggest that stressful environments are not likely to be invaded by most exotic species. However, species, such as H. marinum, that are able to invade these habitats require careful management, especially since these environments often harbor rare species and communities.     

印楝素及印楝杀虫剂的安全性评价研究进展

目前开发的新农药必须具有安全性高、残留低、与环境相容的无公害的农药,从一些植物材料中分离出杀虫活性物质来防治害虫具有较长的历史和优势,从印楝中可以分离出多种对害虫具有活性的化合物如印楝素及等可以有效地控制多种害虫。自印楝素被分离鉴定以来,其广谱而独特的生物活性受到人们的广泛关注,印楝素具有良好的杀虫、拒食、驱虫、昆虫生长调节作用等多种生物活性,在农药、日化等领域都有广泛的应用;由印楝素制备的印楝杀虫制剂对害虫的广谱作用、无残留,可减缓害虫的抗药性,是用于蔬菜、果树、茶叶等作物的理想农药,已成为一种近乎完美的植物源杀虫剂。针对日益严重的农药残留及热点关注的环境安全性等问题,系统总结了印楝素及印楝杀虫剂对人、动物等的毒性及环境安全性评价,将有利于全面评价印楝杀虫剂的安全性及有效地指导印楝杀虫剂开发、生产及推广应用。     

Influence of varying nutrient and pesticide mixtures on abatement efficiency using a vegetated free water surface constructed wetland mesocosm

The nutrient and pesticide abatement efficiency of varying mixtures was examined in a vegetated free water surface constructed wetland. Three different agricultural chemical pollutant mixture conditions were assessed: nutrients only (N and P); pesticides only (atrazine, S-metolachlor and permethrin); and a mixture of nutrients and pesticides. With nutrients only, 672 h nutrient mitigation of 77–91% total phosphorous (TP) and 74–98% total nitrogen (TN) was associated with distance from the injection point and rainfall, whereas with nutrient and pesticide mixtures, 672 h nutrient mitigation of 11–71% TP and 84–98% TN were associated with distance and time. With pesticides only, 672 h pesticide mitigation of 50–99% was associated with distance and time, whereas with nutrients and pesticide mixtures, 672 h pesticide mitigation of 48–99% was associated primarily with distance. Dissipation half-lives were 2–10 times greater for P and 1.5–5 times greater for N when pesticides were present. Pesticide dissipation half-lives showed no clear differences with or without nutrients. While vegetated free water surface constructed wetlands can be effective best management practice tools to trap and abate agricultural run-off during rainfall events, efficiencies can be affected by different types of complex pollutant mixtures and wetland design and implementation should accommodate varying efficiencies.     

浙江省萧山市山一村农村生态良性循环与系统管理研究 Ⅴ.农田病虫害综合防治试验与实施

本项调控工程分两步:1984—1986年在山一村106ha农田开展病虫害综合防治试验示范;1987—1989年将试验成果应用于生产。试验研究取得了明显的经济效益、生态效益和社会效益:农药用量比1983年前减少80.6％,稻谷中农药残留不超标(原有机氯农药666残留超标率31.6％);害虫天敌稻田蜘蛛增加83.3—180.0％,稻虱多索线虫增加2倍;防治农药成本减少53.7％。并制定了《山一村农田农药污染防治技术规程》,其主要内容有:开展病虫害“二查二定”,达到防治指标时用药;制定病虫害防治策略,在防治适期内用药;推广高效低残留农药,遵守农药安全间隔期;推广抗性良种,适施N肥,增施P、K肥;保护利用天敌,发挥天敌自然抑制作用。     

Potential effects of environmental contamination on Yuma Myotis demography and population growth.

Unplanned natural and anthropogenic disasters provide unique opportunities for investigating the influence of perturbations on population vital rates and species recovery times. We investigated the potential effects of a major pesticide spill by comparing annual survival rates using mark-recapture techniques on a riparian bat species, Yuma Myotis (Myotis yumanensis). Demography and population dynamics for most bat species remain poorly understood despite advances in mark-recapture estimation and modeling techniques. We compared survival and population growth rates of two roost populations exposed to a large chemical (metam sodium) spill in the upper Sacramento River in Northern California with two roost populations outside the contaminated area from 1992 to 1996. Hypotheses about long-term effects of the spill on female juvenile and adult survival were tested using an information-theoretic approach (AIC). Working hypotheses included effects of age, chemical spill, and time trend on survival. Female adult survival was higher than female juvenile survival across all sites, suggesting stage-specific mortality risks. Model-averaged estimates of female juvenile survival in the contaminated area (0.50-0.74) were lower than in control roosts (0.60-0.78) for each year in the study, suggesting that the spill may have reduced juvenile survival for several years. Female adult survival (0.73-0.89) did not appear to be strongly affected by the spill during the years of the study. There was an increase in survival for both stage-classes across all populations during the study period, which may have been caused by the end of an extended drought in California in the winter of 1993. The spill-affected population was in decline for the first year of the study as indicated by an estimated growth rate (lambda) < 1, but population growth rates increased during the four-year period.