Summary 1.Competitor, stress-tolerator, ruderal (CSR) theory is a prominent plant functional strategy scheme previously applied to local floras. Globally, the wide geographic and phylogenetic coverage of available values of leaf area (LA), leaf dry matter content (LDMC) and specific leaf area (SLA) (representing, respectively, interspecific variation in plant size and conservative vs. acquisitive resource economics) promises the general application of CSR strategies across biomes, including the tropical forests hosting a large proportion of Earth's diversity. 2. We used trait variation for 3068 tracheophytes (representing 198 families, six continents and 14 biomes) to create a globally calibrated CSR strategy calculator tool and investigate strategy-environment relationships across biomes world-wide. 3. Due to disparity in trait availability globally, co-inertia analysis was used to check correspondence between a 'wide geographic coverage, few traits' data set and a 'restricted coverage, many traits' subset of 371 species for which 14 whole-plant, flowering, seed and leaf traits (including leaf nitrogen content) were available. CSR strategy/environment relationships within biomes were investigated using fourth-corner and RLQ analyses to determine strategy/climate specializations. 4. Strong, significant concordance (RV = 0Á597; P < 0Á0001) was evident between the 14 trait multivariate space and when only LA, LDMC and SLA were used. 5. Biomes such as tropical moist broadleaf forests exhibited strategy convergence (i.e. clustered around a CS/CSR median; C:S:R = 43:42:15%), with CS-selection associated with warm, stable situations (lesser temperature seasonality), with greater annual precipitation and potential evapotranspiration. Other biomes were characterized by strategy divergence: for example, deserts varied between xeromorphic perennials such as Larrea divaricata, classified as S-selected (C:S:R = 1:99:0%) and broadly R-selected annual herbs (e.g. Claytonia perfoliata; R/CR-selected; C:S:R = 21:0:79%). Strategy convergence was evident for several growth habits (e.g. trees) but not others (forbs). 6. The CSR strategies of vascular plants can now be compared quantitatively within and between biomes at the global scale. Through known linkages between underlying leaf traits and growth rates, herbivory and decomposition rates, this method and the strategy-environment relationships it elucidates will help to predict which kinds of species may assemble in response to changes in biogeochemical cycles, climate and land use.
We reviewed the evidence on the role of ants as plant biotic defenses, by conducting meta-analyses for the effects of experimental removal of ants on plant herbivory and fitness with data pooled from 81 studies. Effects reviewed were plant herbivory, herbivore abundance, hemipteran abundance, predator abundance, plant biomass and reproduction in studies where ants were experimentally removed (n = 273 independent comparisons). Ant removal exhibited strong effects on herbivory rates, as plants without ants suffered almost twice as much damage and exhibited 50% more herbivores than plants with ants. Ants also influenced several parameters of plant fitness, as plants without ants suffered a reduction in biomass (-23.7%), leaf production (-51.8%), and reproduction (-24.3%). Effects were much stronger in tropical regions compared to temperate ones. Tropical plants suffered almost threefold higher herbivore damage than plants from temperate regions and exhibited three times more herbivores. Ant removal in tropical plants resulted in a decrease in plant fitness of about 59%, whereas in temperate plants this reduction was not statistically significant. Ant removal effects were also more important in obligate ant-plants (=myrmecophytes) compared to plants exhibiting facultative relationships with hemiptera or those plants with extrafloral nectaries and food bodies. When only tropical plants were considered and the strength of the association between ants and plants taken into account, plants with obligate association with ants exhibited almost four times higher herbivory compared to plants with facultative associations with ants, but similar reductions in plant reproduction. The removal of a single ant species increased plant herbivory by almost three times compared to the removal of several ant species. Altogether, these results suggest that ants do act as plant biotic defenses, but the effects of their presence are more pronounced in tropical systems, especially in myrmecophytic plants.
Major hypotheses on the adaptive significance of insect gall formation are reviewed: nonadaptive, plant protection, mutual benefit, nutrition, microenvironment, and enemy hypotheses. We evaluate the validity of each, and find the first three to be without merit because galls clearly have adaptive features for the insect, but few if any for the plant, and the galler has negative impact on the plant, making the relationship parasitic. Predictions are developed to enable testing of hypotheses, and tests are discussed. Nutrition and microenvironment hypotheses are supported, while the enemy hypothesis remains with several uncertain issues to be resolved. The evolution of the galling habit has followed two pathways, one via mining plant tissues and the other from sedentary external herbivores that then modify plant growth. In each route the sequence of selective factors was probably different, but improved protection from hygrothermal stress and improved nutrition are of pr~mary importance, and protection from enemies probably reinforced the galling habit. KEY WORDS evolution. selection 'f';
We studied the relationship between habitat moisture and gall-forming insect populations. Population sizes for most galling taxa were significantly larger in xeric habitats compared with mesic habitats. Our results indicate that the differential abundance of galling insects in these habitats is due primarily to differential mortality and survivorship. Mortality factors acting upon eight insect galling species (belonging to eight genera and four families) were measured on six species (five genera and five families) of host plants. Survival was significantly higher for galling populations inhabiting xeric habitats compared with mesic habitats. Parasitism was higher in mesic habitats in seven of eight habitats and fungus-induced diseases were higher in five of seven habitats. Mortality due to predation and other (unknown) factors showed no clear trends. Overall, there was a tendency towards lower mortality and consequently higher survival for populations inhabiting xeric habitats. We hypothesize that reduced mortality caused by natural enemies and endophytic fungi has contributed to the speciation and radiation of galling insects in apparently harsh environments.
The plant stress and plant vigor hypotheses (PVH) are two of the most widely recognized hypothesis invoked to explain differential distribution of insect herbivores among their host plants. In both cases, the emphasis is on bottomÁup processes (i.e. host-plant quality), but a recent meta-analytical review of the literature has shown that the plant stress hypothesis might have limited support among insect herbivores. In this study, we conducted a meta-analysis of the effects of plant vigor on insect herbivore abundance and survivorship by reviewing 71 published articles that explicitly tested the PVH and enabled 161 independent comparisons. Z-transform was used as the metric to standardize the results of all independent comparisons. Our quantitative results have shown that Hymenoptera (sawflies) was the most abundant group in the reviewed studies, representing 28.1% of the independent comparisons, followed by Diptera (25.1%) and Homoptera (22.6%). Amongst all the guilds studied, gall-formers were the most representative group (68.0%), whereas leaf-miners and stem-borers were underrepresented (less than 4.0% of the available comparisons). Insect herbivores were significantly more abundant on more vigorous plants (E'' 00.6432, CI 00.7558Á0.7280), but no significant effect was detected on herbivore survivorship. When herbivores were categorized into feeding guilds, effects of plant vigor on herbivore abundance were stronger for sap-suckers, leaf-miners and gall-formers. Our results have shown a strong herbivore preference for more vigorous plants, although our results do not support a preferenceÁperformance linkage.An increased number of studies in plantÁherbivore interactions have recognized that intra-plant heterogeneity affects both the distribution and the performance of herbivorous insects (Schultz 1983, Strong et al. 1984, Crawley and Akhteruzzaman 1988. The plant stress and plant vigor hypotheses are two of the most widely recognized hypotheses invoked to explain heterogeneous distributions of herbivores among their host plants (Williams and Cronin 2004). The plant stress hypothesis (PSH) proposed by White (1984) argues that herbivore abundance is higher on stressed host plants due to an increased availability of nutrients, a decreased concentration of defensive compounds and/or changes in the ratio of nutrients to chemical defenses. Evidence supporting the prediction that moderate stress benefits herbivores due to increased nutritional quality are abundant (McClure 1980, Lewis 1984, Mattson and Haack 1987 and positive relationships between insect herbivory and plant stress have been found for some tree species, crops and herbaceous plants (Mattson and Haack 1987, Heinrichs 1988). Nevertheless, some authors claim that experimental tests of the PSH have generated conflicting results (Bultman and Faeth 1987, Louda and Collinge 1992, Koricheva et al. 1998, De Bruyn et al. 2002, and many authors (Larson 1989, Koricheva et al. 1998) have championed the abandonment of the PSH and the search for alternative hypothes...
One hundred thirty-seven morphotypes of insect galls were found on 73 plant species (47 genera and 30 families) in Serra de São José, in Tiradentes, MG, Brazil. Fabaceae, Myrtaceae, Asteraceae, and Melastomataceae were the plant families that supported most of the galls (49.6% of the total). Galls were mostly found on leaves and stems (66.4% and 25.5%, respectively). Galls were induced by Diptera, Lepidoptera, Coleoptera, Hemiptera (Sternorrhyncha), Hymenoptera, and Thysanoptera. The majority of them (73.7%) were induced by gall midges (Cecidomyiidae: Diptera). Besides the gall inducers, other insects found associated with the galls were parasitoids (Hymenoptera), inquilines (Coleoptera, Lepidoptera, Diptera, and Hemiptera), and predators (Diptera).
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