Xylophagous leafhopppers are common and abundant insects of tropical and subtropical environments and play important ecological roles in these ecosystems. The feeding biology of these insects is unique in terms of their high feeding rates and a digestive physiology that allows them to assimilate amino acids, organic acids, and sugars at approximately 99% efficiency. For those species well studied, fluctuations in plant xylem chemistry and tension appear to determine the diurnal and seasonal use of their host plants. Relatively few species of xylem fluid-feeding leafhoppers are considered important pests in commercial agriculture, as they transmit the bacterial plant pathogen Xylella fastidiosa. X. fastidiosa induces diseases of grapevines, citrus, coffee, almond, alfalfa, stone fruits, landscape ornamentals, and native hardwoods for which there is no cure. Two Xylella diseases, citrus variegated chlorosis (CVC) and Pierce's disease (PD) of grapevines, have emerged as important issues within the past decade. In Brazil, CVC became important in the early 1990s and has now expanded throughout many citrus-growing areas of South America and threatens to spread to North America. The recent establishment of the exotic glassy-winged sharpshooter (Homalodisca coagulata) in California now threatens much of the United States' wine grape, table grape, and almond production. The spread of H. coagulata throughout southern California and the spread of CVC northward from Argentina through Brazil exemplifies the biological risks from exotic species. The occurrence and epidemiology of leafhopper-vectored Xylella diseases are discussed.
Abstract. Enemy‐free space (EFS) was defined by Jeffries & Lawton (1984) as ‘ways of living that reduce or eliminate a species’ vulnerability to one or more species of natural enemies’. EFS has emerged in the literature as a significant niche‐moulding factor. However, the lack of consistency among the empirical studies as to how EFS should be defined, and what hypotheses should be tested in order to evaluate its relative importance, prompted us to review the literature and to propose a working definition that results in a general set of testable hypotheses. To test the relative importance of EFS in structuring the communities of organisms, we propose a set of three falsifiable null hypotheses that must be tested sequentially and rejected. Ho1: The fitness of the organism in an original habit (e.g. on an original host plant) in the presence of natural enemies is equal to the fitness of the organism in that habit in the absence of natural enemies. Acceptance of the alternative hypothesis that the fitness of the organism in the presence of natural enemies is less than in the absence of natural enemies is necessary to demonstrate the importance of natural enemies. Ho2: The fitness of the organism in an alternative habit with natural enemies is equal to the fitness of the organism in the original habit with natural enemies. Acceptance of the alternative hypothesis that the fitness of the organism in the alternative habit with natural enemies is greater than that in the original habit with natural enemies is necessary to demonstrate that the alternative habit provides EFS. Ho3: The fitness of the organism in an alternative habit without natural enemies equals the fitness of the organism in the original habit without natural enemies. Acceptance of the alternative hypothesis that the fitness of the organism in an alternative habit without natural enemies is less than in the original habit without natural enemies is necessary to demonstrate the relative importance of EFS compared with other co‐occurring niche‐moulding factors such as competition or host nutritional quality. We searched the literature and evaluated fifty‐three references (nineteen references to seventeen different terrestrial systems and thirty‐four references to twenty‐four different freshwater systems) to test our hypotheses. Of the forty‐one systems examined, nineteen (46%) tested only for differences in vulnerability of the prey or host species between EFS and non‐EFS options (our Ho2); sixteen (39%) tested for the importance of natural enemies and the effectiveness of the alternative habit in providing EFS (our Ho1 and Ho2); and only ten systems (24%) tested for Ho1, Ho2 and the relative importance of EFS in the system as measured by fitness (our Ho3). Of the systems that tested for EFS, sixteen of nineteen (84%), thirteen of sixteen (81%) and seven of ten (70%) showed evidence in support of the existence of EFS according to hypothesis Ho2 only, hypotheses Ho1 and Ho2, and our three working hypotheses, respectively. These results indicate that very few studies have actually tested for the existence of EFS. Nevertheless, results from this limited number of natural systems suggest that EFS may be important in moulding the niches of arthropods. Because of the large number of claims for EFS in systems where none of the basic hypotheses were investigated, we suggest that authors test for EFS experimentally, be judicious in selecting articles to cite in support of EFS, and exert care in attributing it as a selective force in the evolution of arthropods in specific systems.
Predicting changes in potential habitat for endangered species as a result of global warming requires considering more than future climate conditions; it is also necessary to evaluate biotic associations. Most distribution models predicting species responses to climate change include climate variables and occasionally topographic and edaphic parameters, rarely are biotic interactions included. Here, we incorporate biotic interactions into niche models to predict suitable habitat for species under altered climates. We constructed and evaluated niche models for an endangered butterfly and a threatened bird species, both are habitat specialists restricted to semiarid shrublands of southern California. To incorporate their dependency on shrubs, we first developed climate-based niche models for shrubland vegetation and individual shrub species. We also developed models for the butterfly's larval host plants. Outputs from these models were included in the environmental variable dataset used to create butterfly and bird niche models. For both animal species, abiotic-biotic models outperformed the climate-only model, with climate-only models over-predicting suitable habitat under current climate conditions. We used the climate-only and abiotic-biotic models to calculate amounts of suitable habitat under altered climates and to evaluate species' sensitivities to climate change. We varied temperature ( 1 0.6, 1 1.7, and 1 2.8 1C) and precipitation (50%, 90%, 100%, 110%, and 150%) relative to current climate averages and within ranges predicted by global climate change models. Suitable habitat for each species was reduced at all levels of temperature increase. Both species were sensitive to precipitation changes, particularly increases. Under altered climates, including biotic variables reduced habitat by 68-100% relative to the climate-only model. To design reserve systems conserving sensitive species under global warming, it is important to consider biotic interactions, particularly for habitat specialists and species with strong dependencies on other species.
The glassy-winged sharpshooter (Homalodisca coagulate), an insect that has recently invaded California, and the smoke tree sharpshooter (Homalodisca lacerta) are creating serious new economic problems in both agricultural and ornamental plantings. The greatest threats surround their ability to spread the plantpathogenic bacterium Xylella fastidiosa, which induces Pierce's disease in grapevines, almond leaf scorch disease, and a new disease known as oleander leaf scorch. Because of the potential distribution and large host range of the glassywinged sharpshooter, other diseases caused by different strains of X. fastidiosa may have an even greater impact on the state's agricultural and ornamental landscape industries in the future. Posted 27 June 2000.
Raubenheimer & Simpson (1992) recently discussed the advantages of using analysis of covariance in insect dietary studies as an alternative to the more conventional ratio‐based nutritional indices. We expand on some interpretations of Raubenheimer & Simpson and illustrate our points with examples from published and unpublished data sets. Specifically, we show that an ANCOVA on biomass gain incorporating initial biomass as a covariate provides information not immediately available using the analysis suggested by Raubenheimer & Simpson (ANCOVA on final biomass incorporating initial biomass as a covariate). Second, we show that dietary studies in which the covariate (food consumption) is affected by diet can provide information about the relative importance of preingestive effects (e.g., deterrence) and postingestive effects (e.g., antibiosis) on performance. Cautions about this latter use of ANCOVA are discussed.
Population parameters of the melon aphid, Aphis gossypii Glover, were compared for insects grown on six cultivars of chrysanthemum, Dendranthema grandiflora Tzvelev ('Fontana', 'Iridon', 'Pink Lady', 'Splendor', 'White Diamond', and 'White View Time') fertilized with 3 different levels of nitrogen (80, 160 and 240 mg N/l) and 2 levels of irrigation (high 300 ml and low 210 ml every other day). Fecundity, longevity, and survivorship of aphids were significantly affected by the cultivar treatment, but were not affected by irrigation or fertilizer treatments or by any combination of interactions among cultivar, fertilizer and irrigation. Significantly fewer aphids survived on the cultivar 'Pink Lady' (populations reduced by 20.8%) than any other cultivar examined. Intrinsic rates of increase (r m ), finite rates of increase (R 0 ), and population doubling times (DT) for aphids were different among fertilizer levels and cultivars, but were not different among irrigation levels. There were no significant fertilizer by irrigation interaction effects upon r m , R 0 , or DT. The highest rate of fertilizer applied (240 mg N/liter) adversely affected the intrinsic rate of increase of melon aphids; however, aphid intrinsic rate of increase was weakly correlated with foliar soluble protein levels (r = 0.989, P= 0.0954). Foliar soluble protein levels were not associated with fertilization treatment. Melon aphids perform consistently better on the cultivars 'White Diamond', 'Fontana', and 'Splendor', relative to those aphids growing on 'Iridon', 'Pink Lady', or 'White View Time'. In general, melon aphids performed best on 'White Diamond' and poorest on 'Pink Lady'.
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