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Cicadas have amongst the longest development times and are also amongst the largest insects. Cicadas feed exclusively on xylem fluid, which is nutritionally dilute and difficult to obtain. One possible explanation for their slow development is that poor nutrition limits their growth rate. An analysis of 30 cicada species with known development times is consistent with this hypothesis as species with more equatorial distributions grew more rapidly than those at higher latitudes. A second possible explanation is that prolonged development maximizes net reproductive rate because there is little risk of mortality once early instar cicada nymphs establish feeding sites. Extended development probably allows nymphs to store resources and produce more offspring. Spittlebugs also feed obligately on xylem fluid and grow at similarly slow rates although they attain small adult sizes. Unlike cicadas, spittlebugs do not have steep survivorship curves and have shorter development times. The life histories of cicadas and spittlebugs are therefore consistent with both hypotheses. Cicada development times may be limited if (1) the risk of nymphal mortality equals increased fecundity associated with prolonging development, (2) fluctuating conditions sometimes favour rapid development times, or (3) host plant quality changes over time and penalizes nymphs that cannot relocate feeding sites.

Plants are facing increased risk of heat stress with global climate change. Reproductive tissues are particularly sensitive which can result in lower plant fitness. Floral shading and closure are possible mechanisms to limit heat stress although most previous work on petal orientation has considered adaptations to raise temperatures. We hypothesized that floral shading could reduce temperature and increase reproductive success. Methods We measured floral temperatures of four species that exhibited intraspecific variation in flower closure (Opuntia ficus-indica, Oenothera elata, Convolvulus arvensis, and Romneya coulteri). We also wired newly opened R. coulteri flowers so that they were either permanently open, permanently closed, or unencumbered controls. Results Individual flowers of all four species that shaded their pistils experienced temperatures 3 - 8

Communication occurs when a sender emits a cue perceived by a receiver that changes the receiver's behavior. Plants perceive information regarding light, water, other nutrients, touch, herbivores, pathogens, mycorrhizae, and nitrogen-fixing bacteria. Plants also emit cues perceived by other plants, beneficial microbes, herbivores, enemies of herbivores, pollinators, and seed dispersers. Individuals responding to light cues experienced increased fitness. Evidence for benefits of responding to cues involving herbivores and pathogens is more limited. The benefits of emitting cues are also less clear, particularly for plant–plant communication. Reliance on multiple or dosage-dependent cues can reduce inappropriate responses, and plants often remember past cues. Plants have multiple needs and prioritize conflicting cues such that the risk of abiotic stress is treated as greater than that of shading, which is in turn treated as greater than that of consumption. Plants can distinguish self from nonself and kin from strangers. They can identify the species of competitor or consumer and respond appropriately. Cues involving mutualists often contain highly specific information.

PREMISE Ecologists have an incomplete understanding of the factors that select for deciduous, evergreen, and marcescent leaf habits. Evergreens have more opportunities for photosynthesis but may experience costs when abiotic conditions are unfavorable such as during ice and windstorms. METHODS We documented branch loss for species of oaks (Quercus spp.) in a common garden in California during an unusual windstorm. RESULTS Branches of marcescent trees were more likely to break during the storm, and this pattern had a negligible phylogenetic signature. Branches of evergreen and marcescent species were mostly alive before breaking, which likely accrued a fitness cost, while those of deciduous species were mostly already dead. One explanation for the overrepresentation of broken branches from marcescent species is that their petioles are inflexible and have greater wind loading compared to the flexible petioles of evergreen leaves and the leafless condition of deciduous branches. CONCLUSIONS These results suggest that branch loss during unusual winter storms may be an important cost of a marcescent leaf habit.

The interaction between endogenous dynamics and exogenous environmental variation is central to population dynamics. Although investigations into the effects of changing mean climate are widespread, changing patterns of variation in environmental forcing also affect dynamics in complex ways. Using wavelet and time series analyses, we identify a regime shift in the dynamics of a moth species in California from shorter to longer period oscillations over a 34-year census, and contemporaneous changes in regional precipitation dynamics. Simulations support the hypothesis that shifting precipitation dynamics drove changes in moth dynamics, possibly due to stochastic resonance with delayed density-dependence. The observed shift in climate dynamics and the interaction with endogenous dynamics mean that predicting future population dynamics will require information on both climatic shifts and their interaction with endogenous density-dependence, a combination that is rarely available. Consequently, models based on historical data may be unable to predict future population dynamics.

Plants can retain either physiologically active green leaves or inactive brown leaves over winter. Research has suggested that leaf retention incurs a cost due to higher herbivore load in the following year; however, no distinction has, thus, far been made between retention of green and brown leaves. We surveyed the over-winter retention of physiologically active green and inactive brown leaves of valley oaks (Quercus lobata) and examined their relationship with the diversity and density of 15 gall-making cynipid wasp species. Cynipid diversity in the spring was 8.2-fold greater on larger trees with more green leaf retention. Brown leaf retention was not associated with spring cynipid diversity, but was related to a substantial 20-fold reduction in spring cynipid densities on large trees. Retention of either leaf type was generally a poor predictor of summer cynipid diversity and density. Overall, green leaf retention better explained cynipid diversity, but brown leaf retention better explained cynipid densities. These differing effects may be explained by the fact that green leaves provide a common cue used by herbivores to find a suitable host. Retained brown leaves, however, may be an ecological trap for over-winter gall wasps that normally drop to the ground.

Vehicle pollution is a pervasive aspect of anthropogenic change across rural and urban habitats. The most common emissions are carbon- or nitrogen-based pollutants that may impact diverse interactions between plants and insect herbivores. However, the effects of vehicle pollution on plant-insect interactions are poorly understood.Here, we combine a city-wide experiment across the Sacramento Metropolitan Area and a laboratory experiment to determine how vehicle emissions affect insect herbivory and leaf nutritional quality.We demonstrate that leaf damage to a native oak species (Quercus lobata) commonly planted across the western US is substantially elevated on trees exposed to vehicle emissions. In the laboratory, caterpillars preferred leaves from highway-adjacent trees and performed better on leaves from those same trees.Synthesis and applications. Together, our studies demonstrate that the heterogeneity in vehicle emissions across cities may explain highly variable patterns of insect herbivory on street trees. Our results also indicate that trees next to highways are particularly vulnerable to multiple stressors, including insect damage. To combat these effects, urban foresters may consider installing trees that are less susceptible to insect herbivory along heavily traveled roadways.

Dispersal is a key driver of spatial population dynamics. Dispersal behaviour may be shaped by many factors, such as mate-finding, the spatial distribution of resources, or wind and currents, yet most models of spatial dynamics assume random dispersal. We examined the spatial dynamics of a day-flying moth species ( Arctia virginalis ) that forms mating aggregations on hilltops (hilltopping) based on long-term adult and larval population censuses. Using time-series models, we compared spatial population dynamics resulting from empirically founded hilltop-based connectivity indices and modelled the interactive effects of temperature, precipitation and density dependence. Model comparisons supported hilltop-based connectivity metrics including hilltop elevation over random connectivity, suggesting an effect of hilltopping behaviour on dynamics. We also found strong interactive effects of temperature and precipitation on dynamics. Simulations based on fitted time-series models showed lower patch occupancy and regional synchrony, and higher colonization and extinction rates when hilltopping was included, with potential implications for the probability of persistence of the patch network. Overall, our results show the potential for dispersal behaviour to have important effects on spatial population dynamics and persistence, and we advocate the inclusion of such non-random dispersal in metapopulation models.

Infectious disease is an important potential driver of population cycles, but this must occur through delayed density-dependent infection and resulting fitness effects. Delayed density-dependent infection by baculoviruses can be caused by environmental persistence of viral occlusion bodies, which can be influenced by environmental factors. In particular, ultraviolet radiation is potentially important in reducing the environmental persistence of viruses by inactivating viral occlusion bodies.Delayed density-dependent viral infection has rarely been observed empirically at the population level although theory predicts that it is necessary for these pathogens to drive population cycles. Similarly, field studies have not examined the potential effects of ultraviolet radiation on viral infection rates in natural animal populations. We tested if viral infection is delayed density-dependent with the potential to drive cyclic dynamics and if ultraviolet radiation influences viral infection levels.We censused 18 Ranchman’s tiger moth (Arctia virginalis) populations across nearly 9° of latitude over two years and quantified the effects of direct and delayed density and ultraviolet radiation on baculovirus infection rates, infection severity, and survival to adulthood. Caterpillars were collected from each population in the field and reared in the laboratory. Baculovirus has not previously been described infecting Arctia virginalis, and we used genetic methods to confirm the identity of the virus.We found that infection rate, infection severity, and survival to adulthood exhibited delayed density-dependence. Ultraviolet radiation in the previous summer decreased infection severity, and increased survival probability of the virus. Structural equation modelling indicated that the effect of lagged density on moth survival was mediated through infection rate and infection severity, and was 2.5 fold stronger than the effect of ultraviolet radiation on survival through infection severity. We successfully amplified polh, lef-8, and lef-9 viral genes from caterpillar samples, and BLAST search results confirmed that the virus was a nucleopolyhedrovirus.Our findings provide clear evidence that delayed density dependence can arise through viral infection rate and severity in insects, which supports the role of viral disease as a potential mechanism, among others, that may drive insect population cycles. Furthermore, our findings support predictions that ultraviolet radiation can modify viral disease dynamics in insect populations, most likely through attenuating viral persistence in the environment.

Animal biologists have recently focused on individual variation in behavioral traits and have found that individuals of many species have personalities. These are defined as consistent intraspecific differences in behaviors that are repeatable across different situations and stable over time. When animals sense danger, some individuals will alert neighbors with alarm calls and both calling and responding vary consistently among individuals. Plants, including sagebrush, emit volatile cues when they are attacked by herbivores and neighbors perceive these cues and reduce their own damage. We experimentally transferred volatiles between pairs of sagebrush plants to evaluate whether individuals showed consistent variation in their effectiveness as emitters and as receivers of cues, measured in terms of reduced herbivore damage. We found that 64% of the variance in chewing damage to branches over the growing season was attributable to the identity of the individual receiving the cues. This variation could have been caused by inherent differences in the plants as well as by differences in the environments where they grew and their histories. We found that 5% of the variance in chewing damage was attributable to the identity of the emitter that provided the cue. This fraction of variation was statistically significant and could not be attributed to the environmental conditions of the receiver. Effective receivers were also relatively effective emitters, indicating consistency across different situations. Pairs of receivers and emitters that were effective communicators in 2018 were again relatively effective in 2019, indicating consistency over time. These results suggest that plants have repeatable individual personalities with respect to alarm calls.

Plant-to-plant volatile-mediated communication and subsequent induced resistance to insect herbivores is common. Less clear is the adaptive significance of these interactions; what selective mechanisms favour plant communication and what conditions allow individuals to benefit by both emitting and responding to cues? We explored the predictions of two non-exclusive hypotheses to explain why plants might emit cues, the kin selection hypothesis (KSH) and the mutual benefit hypothesis (MBH). We examined 15 populations of sagebrush that experience a range of naturally occurring herbivory along a 300 km latitudinal transect. As predicted by the KSH, we found several uncommon chemotypes with some chemotypes occurring only within a single population. Consistent with the MBH, chemotypic diversity was negatively correlated with herbivore pressure; sites with higher levels of herbivory were associated with a few common cues broadly recognized by most individuals. These cues varied among different populations. Our results are similar to those reported for anti-predator signalling in vertebrates.

Plant induced defenses may benefit plants by increasing cannibalism among insect herbivores. However, the general efficacy of plant defenses that promote cannibalism remains unclear. Using a generalist Lepidopteran herbivore (Helicoverpa zea), we examined whether plant induced defenses in Solanum lycopersicum increased cannibalism among H. zea and whether defense-mediated cannibalism benefits both the plant and the cannibal. In a separate experiment, we also examined whether defense-mediated cannibalism has effects on H. zea herbivory that are comparable to the effects of pathogenic virus of H. zea (HzSNPV) and whether defense-mediated cannibalism modified pathogen efficacy. We found that both plant defenses and cannibalism decreased herbivory: H. zea consumed less plant material if plants were induced, if dead conspecifics were provided, or both. Cannibalism increased cannibal growth rate: cannibals effectively overcome the costs of plant defenses by eating conspecifics. Inoculating half of H. zea with virus strongly reduced caterpillar survival. Cannibalism occurred sooner among virus-inoculated groups of H. zea, and all caterpillars in virus-inoculated treatments died before the end of the 7-day experiment. Although the rise in mortality caused by HzSNPV occurred more rapidly than the rise in mortality due to defense-mediated cannibalism, overall H. zea mortality at the end of the experiment was equal among virus-inoculated and induced-defense groups. Defense-mediated cannibalism and viral inoculation equally reduced herbivory on S. lycopersicum. Our results provide evidence that defense-mediated increases in cannibalism can be as effective as other forms of classic herbivore population regulation, and that both viral pathogens and defense-induced cannibalism can have significant benefits for plants.

Apart from model organisms, 13- and 17-year periodical cicadas (Hemiptera: Cicadidae: Magicicada) are among the most studied insects in evolution and ecology. They are attractive subjects because they predictably emerge in large numbers; have a complex biogeography shaped by both spatial and temporal isolation; and include three largely sympatric, parallel species groups that are, in a sense, evolutionary replicates. Magicicada are also relatively easy to capture and manipulate, and their spectacular, synchronized mass emergences facilitate outreach and citizen science opportunities. Since the last major review, studies of Magicicada have revealed insights into reproductive character displacement and the nature of species boundaries, provided additional examples of allochronic speciation, found evidence for repeated and parallel (but noncontemporaneous) evolution of 13- and 17-year life cycles, quantified the amount and direction of gene flow through time, revealed phylogeographic patterning resulting from paleoclimate change, examined the timing of juvenile development, and created hypotheses for the evolution of life-cycle control and the future effects of climate changeon Magicicada life cycles. New ecological studies have supported and questioned the role of prime numbers in Magicicada ecology and evolution, found bidirectional shifts in population size over generations, quantified the contribution of Magicicada to nutrient flow in forest ecosystems, and examined behavioral and biochemical interactions between Magicicada and their fungal parasites and bacterial endosymbionts.

Background and Aims Pendulous flowers (due to a flexible pedicel) are a common, convergent trait of hummingbird-pollinated flowers. However, the role of flexible pedicels remains uncertain despite several functional hypotheses. Here we present and test the ‘lever action hypothesis’: flexible pedicels allow pendulous flowers to move upwards from all sides, pushing the stigma and anthers against the underside of the feeding hummingbird regardless of which nectary is being visited. Methods To test whether this lever action increased pollination success, we wired emasculated flowers of serpentine columbine, Aquilegia eximia, to prevent levering and compared pollination success of immobilized flowers with emasculated unwired and wire controls. Key Results Seed set was significantly lower in wire-immobilized flowers than unwired control and wire control flowers. Video analysis of visits to wire-immobilized and unwired flowers demonstrated that birds contacted the stigmas and anthers of immobilized flowers less often than those of flowers with flexible pedicels. Conclusions We conclude that flexible pedicels permit the levering of reproductive structures onto a hovering bird. Hummingbirds, as uniquely large, hovering pollinators, differ from flies or bees which are too small to cause levering of flowers while hovering. Thus, flexible pedicels may be an adaptation to hummingbird pollination, in particular due to hummingbird size. We further speculate that this mechanism is effective only in radially symmetric flowers; in contrast, zygomorphic hummingbird-pollinated flowers are usually more or less horizontally oriented rather than having pendulous flowers and flexible pedicels.

Patch-based population models predominately focus on factors that affect regional processes namely, patch size and connectivity, as the primary drivers explaining patch occupancy. This trend persists despite the recognition that patch quality can strongly influence population demography at the local scale. The quality of patches is often temporally variable and influenced by abiotic conditions. However, few studies have explicitly investigated how climatic variables influence the spatial and temporal dynamics of spatially-structured populations either directly or indirectly through changes in patch quality. Using a 10-year census of a spatially-structured population of an outbreaking caterpillar, we determined the relative importance of patch quality (determined demographically), connectivity, precipitation, and their interactive effects on patch abundance, occupancy, colonization, and extinction. We generated a series of statistical models and performed comparisons using Akaike’s information criterion. We subsequently used likelihood ratio tests to determine the influence of each parameter on model fit. Patch quality and precipitation were the strongest predictors of the observed dynamics. We found that the dynamics of the spatially-structured population of Arctia virginalis were strongly influenced by precipitation: all patches had a higher probability of occupancy, contained higher abundances of caterpillars, and experienced fewer extinctions following wet winters compared to years following droughts. These findings suggest that precipitation may act to influence the strength of heterogeneity of patch quality. This work demonstrates that patch-based models that do not include local and climatic factors may produce poor predictions under future climatic regimes.

Climate change can affect biotic interactions, and the impacts of climate on biotic interactions may vary across climate gradients. Climate affects biotic interactions through multiple drivers, although few studies have investigated multiple climate drivers in experiments. We examined the effects of experimental watering, warming, and predator access on leaf water content and herbivory rates of woolly bear caterpillars (Arctia virginalis) on a native perennial plant, pacific silverweed (Argentina anserina ssp. pacifica), at two sites across a gradient of precipitation in coastal California. Based on theory, we predicted that watering should increase herbivory at the drier end of the gradient, predation should decrease herbivory, and watering and warming should have positive interacting effects on herbivory. Consistent with our predictions, we found that watering only increased herbivory under drier conditions. However, watering increased leaf water content at both wetter and drier sites. Warming increased herbivory irrespective of local climate and did not interact with watering. Predation did not affect herbivory rates. Given predictions that the study locales will become warmer and drier with climate change, our results suggest that the effects of future warming and drying on herbivory may counteract each other in drier regions of the range of Argentina anserina. Our findings suggest a useful role for range‐limit theory and the stress‐gradient hypothesis in predicting climate change effects on herbivory across stress gradients. Specifically, if climate change decreases stress, herbivory may increase, and vice versa for increasing stress. In addition, our work supports previous suggestions that multiple climate drivers are likely to have dampening effects on biotic interactions due to effects in different directions, though this is context‐dependent., Multiple interacting climate drivers are expected to affect ecological interactions and to vary across climate gradients. In this study, we manipulate temperature and moisture a two sites across a gradient of precipitation and find that warming increases herbivory at both sites, but watering only increases herbivory at the drier site. These findings are consistent with predictions drawn from range‐limit theory and the stress‐gradient hypothesis.