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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

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.

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.

Fire suppression has increased fuel load and the risk of catastrophic wildfire in forest and woodland ecosystems across the Western United States. In an effort to reduce fuel load and restore historical structure and function, land managers have implemented fuel reduction treatments on millions of acres. Reducing fuel loads protects people, structures, and in some cases, improves ecosystem health. However, the ecological risks of soil surface disturbance related to fuel reduction strategies, and subsequent soil erosion, may be significant in some cases. Here, we examined the effects of common fuel reduction strategies (mechanical mastication and two techniques for prescribed burning) on wind and water erosion in two upland pinon-juniper woodlands in SE Utah over 2 years. We also tested the impact of broadcast seeding coupled with fuel reduction as a way to mitigate erosional soil loss. Finally, we analyzed biotic and abiotic predictor variables to evaluate important drivers of soil erosion following fuel treatments. We found that both techniques for prescribed burning—pile burning and broadcast burning—increased wind-related sediment fluxes by an average of 11-fold and 58-fold, respectively. Mastication did not increase wind-related losses over untreated controls. Erosional fluxes measured at silt fences, that captured both wind- and water-driven sediments, followed similar trends with moderate increases from pile burning (fivefold) but larger increases from broadcast burning (17-fold). Seeding did not affect erosion rates. Our results suggest that prescribed fire significantly increases soil erosion in fuel-treated pinon-juniper woodlands and may be a degradation pathway when implementing treatments.

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.

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.

Insect declines have been reported worldwide, although the particular causes of the declines may be complex and are poorly understood. Meadow spittlebugs were one of the most abundant insects in the coastal prairie along the California coast 40 yr ago but have largely disappeared. Evidence links this decline to changing climatic conditions, which have reduced survival of eggs and neonates. We identified several refugia where meadow spittlebug populations have persisted amidst unfavorable conditions. Protection from desiccating winds was the common attribute of these refugia. Following a wet year, adult meadow spittlebugs were able to disperse from one refuge that we studied to recolonize coastal prairie habitats, although populations declined over the next two drier years. Because of their previous high abundance, loss of meadow spittlebugs is likely to affect the functioning of this widespread habitat, including energy transfer, their host plants, and their predators. In addition, meadow spittlebugs are unusual in having been the subject of extensive physiological and long‐term ecological data, so they can serve as a bellwether species, indicating the effects of climate change.

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.

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.

Induced plant responses to herbivory are common, and we have learned a lot about the mechanisms of induced resistance and their effects on herbivore performance. We know less about their effects on herbivore behaviour and especially on spatial patterns of damage. Theoretical models predict that induced responses can cause patterns of damage to become aggregated, random or even. A recent model predicted that informed herbivore movement coupled with communication between plants would make damage more even within individual plants. We tested these predictions in the field using a specialist beetle Trirhabda pilosa that feeds on sagebrush Artemisia tridentata. Both the beetle and the plant are well-documented to respond to damage-induced volatile cues. Beetle larvae were more likely to move from damaged leaves and leaves that had been exposed to volatiles from nearby damaged leaves compared to undamaged control leaves. Previous laboratory results indicated that beetles were more likely to choose undamaged leaves compared to damaged leaves or those exposed to volatile cues of damage. A comparison of damage patterns early in the season and after completion of beetle feeding revealed that variance in damage among branches decreased as the season progressed; that is, damage became more evenly distributed among the branches within a plant. Larvae damaged many leaves on a plant but removed relatively little tissue from each leaf. Herbivore movement and the spatial patterns of damage that it creates can be important in determining effects on plant fitness and other population processes. Dispersion of damage deserves more consideration in plant-herbivore studies.

Plants respond to damage by herbivores or to reliable cues of damage by changing in ways that provide greater resistance and increase their fitness. Sagebrush has been a model system for understanding induced resistance, although resistance in this system is commonly assessed by quantifying damage at the end of the season; this measure is slow and lacks accuracy and interpretability because so many other factors also affect levels of damage. Herbivore choice offers a potentially rapid and accurate alternative assay of induced resistance. Here we evaluate whether feeding behavior of a specialized Chrysomelid beetle, Trirhabda pilosa, could be used to assay induced changes in plant quality. Beetle larvae were offered the choice between two leaves in Petri dish arenas. We found that beetles avoided leaves that were naturally damaged by herbivores, experimentally clipped with scissors, and exposed to the volatiles from naturally or experimentally damaged neighboring leaves compared to control leaves. Experiments varied the source of the damage, the duration of the feeding test, and how damage was measured, still, beetles consistently preferred uninduced controls by a 2:1 ratio over leaves exposed to cues of damage. These results suggest that behavioral assays using T. pilosa larvae can be used to rapidly evaluate induced resistance in this system. More generally, movement and feeding behaviors of herbivores are an important and underappreciated component of induced plant responses.

‘Hilltopping’ is a common mate-locating behaviour exhibited by numerous insect taxa; individuals aggregate on summits, ridges and other topographic features, and thereby increase their likelihood of mating. Recently, hilltopping has gained interest as a model system to study nonrandom dispersal. We tested four predictions from the hilltopping literature regarding individual movement behaviour and the resulting spatial distribution of summit aggregations. Through observations and capture–mark–recapture studies using the day-flying tiger moth, Arctia (formerly Platyprepia ) virginalis , we found evidence for all predictions. The highest densities of moths were associated with a few, high-elevation summits and were recaptured over multiple days. No individuals were found to move between summit aggregations and mated females had shorter residency times than males. We discuss our results in the context of the predictions, the behaviour of other hilltopping species, implications for population structure and spatial population dynamics.

© 2017 by the Ecological Society of America Climate is widely recognized as an important factor that affects temporal and spatial patterns of occurrence and abundance of herbivorous insects, although the ecological mechanisms responsible are poorly understood. We found that precipitation and standing water were positively correlated with locations and years of high abundance of caterpillars of the ranchman's tiger moth, Platyprepia virginalis. We analyzed 30 years of survey data and found that the number of large rainfall events was a better predictor of caterpillar abundance than total annual accumulation. We considered three ecological mechanisms that could drive this relationship and conducted observations and manipulative experiments to evaluate these mechanisms. (1) Rainfall facilitates more plant growth, although we found no evidence that increased food quality or quantity was causing the positive association between precipitation and caterpillar abundance. (2) Large rainfall events cause predatory ground-nesting ants to be less abundant and we found that the number of ants that recruited to local sites was negatively associated with survival and abundance of caterpillars. (3) We found that litter from wet sites provided a refuge from ant predation; litter from wet sites was not beneficial to caterpillars in the absence of ants. Both abiotic factors (precipitation) and biotic factors (predatory ants) affected the temporal and spatial abundance of caterpillars directly and interactively. Climate models predict that rainfall will become more variable, suggesting that populations of this caterpillar may also become more variable in the future.

Habitat‐forming species provide refuges for a variety of associating species; these refuges may mediate interactions between species differently depending on the functional traits of the habitat‐forming species. We investigated refuge provisioning by plants with different functional traits for dragonfly and damselfly (Odonata: Anisoptera and Zygoptera) nymphs emerging from water bodies to molt into their adult stage. During this period, nymphs experience high levels of predation by birds. On the shores of a small pond, plants with mechanical defenses (e.g., thorns and prickles) and high structural complexity had higher abundances of odonate exuviae than nearby plants which lacked mechanical defenses and exhibited low structural complexity. To disentangle the relative effects of these two potentially important functional traits on nymph emergence‐site preference and survival, we conducted two fully crossed factorial field experiments using artificial plants. Nymphs showed a strong preference for artificial plants with high structural complexity and to a lesser extent, mechanical defenses. Both functional traits increased nymph survival but through different mechanisms. We suggest that future investigations attempt to experimentally separate the elements contributing to structural complexity to elucidate the mechanistic underpinnings of refuge provisioning.

Seed mucilage, a coating on seeds or fruit that becomes slimy and sticky when wet, has evolved convergently many times across plants. One common consequence of having seed mucilage is that sand and dirt particles stick to wet seeds and remain tightly bound to the seed surface after the mucilage dries. Here, we test the hypothesis that a mucilage-bound sand coating protects the seed from seed predators; either as a physical barrier or by reducing apparency of the seed (i.e., camouflage). We experimentally manipulated the sand coating on seeds of 53 plant species of 13 families and assayed the defensive benefit of the sand coating in feeding "depots" near harvester ant nests in California's Central Valley. Consistent with a defensive function, sand coating reduced ant predation on seeds in 48 of the 53 species examined. To test whether this striking benefit was due to reduced apparency, we conducted an addition experiment using flax seeds in which we factorially manipulated the color of both the background substrate and the sand coating, creating visually apparent and unapparent seeds. Our results did not support the reduced apparency hypothesis; seeds coated in background-matched sand were removed at the same rate as seeds coated in unmatched sand. The defensive benefit of a sand coating was not well-predicted by seed mass, entrapped sand mass, or sand mass scaled by seed mass. Together, our results demonstrate that seed mucilage is a phylogenetically widespread and effective seed defensive trait and point to the physical barrier, not reduced apparency, as a mechanism.

Author(s): Wetzel, W; Kharouba, H; Robinson, M; Holyoak, M; Karban, R | Abstract: SADIE (Spatial Analysis by Distance Indices) is designed specifically to quantify patterns in spatially-referenced count-based data. It was developed for dealing with data that can be considered ‘patchy’. Such distributions are commonly found, for example, in insect populations where discrete patches of individuals are often evident. The distributions of such populations have ‘hard edges’, with patches and gaps occurring spatially. In these cases variance of abundance does not vary smoothly, but discontinuously. In this paper we outline the use of SADIE and provide free access to the SADIE software suite, establishing Rethinking Ecology as its permanent home. Finally, we review the use of SADIE and demonstrate its use in a wide variety of sub-disciplines within the general field of ecology.

© 2018, Springer Nature B.V. Grasses frequently have unidirectional hairs, prickles, and spines; these leaf features have been hypothesized to move herbivores and their chewing damage away from grass meristems, which are located basally. Observations of chewing damage to two grasses, Andropogon virginicus and Phragmites australis, were consistent with this hypothesis as leaf tips received 10 × and 2 × more damage than bases. Grasshoppers were no more likely to land on leaf tips than bases although they oriented towards the tips after landing. Leaves of A. virginicus that were damaged by chewing herbivores had fewer spines than leaves on the same or neighboring plants that lacked damage. This suggests that herbivores chose less spiny leaves. At a larger spatial scale, plants in neighborhoods favorable for grasshoppers had more spines than plants in less favorable neighborhoods. We found no evidence that marginal spines allowed leaves to shed water more rapidly, a potential alternative benefit. The density of spines on new leaves increased following cues of damage. A. virginicus leaves produced after an adjacent leaf had been clipped with scissors had 13% more spines than new leaves on unclipped plants. Clipping with scissors failed to increase spine density for new P. australis leaves although experimental chewing by caterpillars led to the production of new leaves with 24% more spines than controls. Unchewed new leaves within 20 cm of a chewed neighbor had 13% more spines than controls. Grasses are capable of responding to cues of tissue damage to their own and neighboring leaves, potentially reducing herbivory to meristems.

Although many studies have investigated plant growth in the context of episodic herbivory and pressed resource availability, relatively few have examined how plant growth is affected by pulsed resources and chronic herbivory. Periodical cicada (Magicicada spp.) adults represent a pulsed detrital subsidy that fertilizes plants, and live cicada nymphs are long-lived root-feeding herbivores. Previous studies of cicada herbivory effects have been inconclusive, and previous studies of cicada-mediated fertilization did not examine effects on trees, or on a multiyear timescale. Here, we describe the results of a 3-yr experiment that factorially manipulated the presence and absence of cicada fertilization and herbivory in a population of 100 American sycamore (Platanus occidentalis) trees. We found that cicada fertilization strongly increased tree growth in the year of emergence, creating differences in tree size that persisted at least 2yr later. By comparison, we did not detect reductions in tree growth associated with cicada herbivory in any year of this experiment. However, cicada herbivory reduced the densities of, and damage from, other aboveground herbivores. These results suggest that cicadas affect the size structure of forests over multiple years, and raise questions about how cicada-mediated fertilization and herbivory will affect tree growth over longer timescales.

Many plants engage in protective mutualisms, offering resources such as extrafloral nectar and shelters to predatory arthropods in exchange for protection against herbivores. Recent work indicates that sticky plants catch small insects and provide this carrion to predators who defend the plants against herbivores. In this study, we investigated whether wild tobacco, Nicotiana attenuata, fits this sticky plant defense syndrome that has been described for other sticky plants. We developed a bioassay for stickiness involving the number of flies that adhered to flowers, the stickiest tissues. In surveys conducted over three field seasons at four sites, we found that the number of carrion that adhered to a plant was positively correlated with the number of predators that we observed foraging over its surfaces. The number of predators was positively correlated with the number of seed capsules that the plant produced, a measure of lifetime female reproductive success. Structural equationmodeling indicated strong support for the causal path linking carrion numbers to predator numbers to capsule production. We investigated whether stickiness was an inducible trait and examined two potential cues. We found that experimental clipping of rosette leaves induced greater stickiness, although clipping of neighboring sagebrush leaves did not. Damage to leaf tissue is likely to be a more reliable predictor of risk than is damage to a neighboring plant. The sticky plant defense syndrome is a widespread protective mutualism; its strength and ecological relevance can adjust as risk of herbivory changes.

Sand entrapment on plant surfaces, termed psammophory or sand armor, is a phylogenetically and geographically widespread trait. The functional significance of this phenomenon has been poorly investigated. Sand and soil are nonnutritive and difficult for herbivores to process, as well as visually identical to the background. We experimentally investigated whether this sand coating physically protected the plant from herbivores or increased crypsis (e.g., decreased apparency to herbivores). We tested the former hypothesis by removing entrapped sand from stems, petioles, and leaves of the sand verbena Abronia latifolia and by supplementing natural sand levels in the honeyscented pincushion plant Navarretia mellita. Consistent with a physical defensive function, leaves with sand present or supplemented suffered less chewing herbivory than those with sand removed or left as is. To test a possible crypsis effect, we coated some sand verbena stems with green sand, matching the stem color, as well as others with brown sand to match the background color. Both suffered less chewing herbivory than controls with no sand and herbivory did not significantly differ between the colors, suggesting crypsis was not the driving resistance mechanism. Strong tests of plant apparency are rare; this experimental approach may be possible in other systems and represents one of few manipulative tests of this long-standing hypothesis.

The risk of consumption is a pervasive aspect of ecology and recent work has focused on synthesis of consumer-resource interactions (e.g., enemy-victim ecology). Despite this, theories pertaining to the timing and magnitude of defenses in animals and plants have largely developed independently. However, both animals and plants share the common dilemma of uncertainty of attack, can gather information from the environment to predict future attacks and alter their defensive investment accordingly. Here, we present a novel, unifying framework based on the way an organism's ability to defend itself during an attack can shape their pre-attack investment in defense. This framework provides a useful perspective on the nature of information use and variation in defensive investment across the sequence of attack-related events, both within and among species. It predicts that organisms with greater proportional fitness loss if attacked will gather and respond to risk information earlier in the attack sequence, while those that have lower proportional fitness loss may wait until attack is underway. This framework offers a common platform to compare and discuss consumer effects and provides novel insights into the way risk information can propagate through populations, communities, and ecosystems.

Grasses are major agricultural products worldwide and they are critical to ecosystem function in many terrestrial habitats. Despite their global importance, we know relatively little about their defenses against herbivory. Grasses tend to be tolerant of leaf loss because their valuable meristems are located underground, out of reach for above ground herbivores. Many grasses have unidirectional leaf hairs, prickles, and spines that make moving up the leaf blade easy, but make moving down, toward the meristem, difficult. We tested the hypothesis that unidirectional grass hairs direct small arthropod herbivores away from the meristems. In a field survey of the distribution of herbivore damage, we found that leaf tips received five times more damage than leaf bases for Avena barbata. Early-instar grasshoppers fed three times as often on leaf tops as on leaf bases of pubescent individuals in a common garden laboratory experiment. This effect was not observed for glabrous individuals where grasshoppers damaged leaf bases as often as leaf tops. A common generalist caterpillar, Heliothus virescens, was more than twice as likely to turn in the direction of the hairs, away from the meristems, when it encountered pubescent leaves of A. barbata. However, larger caterpillars of the generalist feeder Arctia virginalis showed no directional bias when they encountered pubescent leaves. In common garden experiments, selection on pubescence was weak and inconsistent over space and time. Under some circumstances, individuals of A. barbata with pubescent leaves were more likely to produce seeds than were individuals with fewer hairs. The surveys, behavioral experiments with small insects, and estimates of lifetime reproduction all support the hypothesis that unidirectional leaf hairs on A. barbata, and perhaps other grasses, serve as an unstudied defense that direct small herbivores away from the meristems.

© 2018 by the Ecological Society of America Climate change can have strong effects on species interactions and community structure. Temperature-dependent effects on predator–prey interactions are a major mechanism through which these effects occur. To understand the net effects of predator attack rates and dynamic windows of prey vulnerability, we examined the impacts of temperature on the interaction of a caterpillar (Arctia virginalis) and its ant predator (Formica lasioides). We conducted field experiments to examine attack rates on caterpillars relative to temperature, ant abundance, and body size, and laboratory experiments to determine the effects of temperature on caterpillar growth. We modeled temperature-dependent survival based on the integrated effects of temperature-dependent growth and temperature- and size-dependent predation. Attack rates on caterpillars increased with warming and ant recruitment, but decreased with caterpillar size. Caterpillar growth rates increased with temperature, narrowing the window of vulnerability. The model predicted that net caterpillar survival would decrease with temperature, suggesting that A. virginalis populations could be depressed with future climate warming. Theoretical work suggests that the net outcome of predator–prey interactions with increasing temperature depends on the respective responses of interacting species in terms of velocity across space, whereas the present study suggests the importance of effects of temperature on prey window of vulnerability, or “velocity” across time.

Entomopathogenic nematodes (EPNs) can kill and regulate populations of soil-inhabiting insects, but studies evaluating these interactions in native ecosystems are rare. The objective of this study was to examine the effects of EPNs on a non-agricultural caterpillar, Platyprepia virginalis (Boisduval) (Lepidoptera: Arctiidae), under natural conditions. Platyprepia virginalis caterpillars live in litter on the soil surface feeding beneath bush lupine during summer, autumn, and winter. Initial laboratory assays revealed that the caterpillars were vulnerable to at least two species of EPNs with which they co-occur in the coastal prairie in northern California (USA). In contrast to laboratory assays, caterpillars survived exposure to prairie soil containing EPNs under natural conditions in field assays. To better understand the divergence between laboratory and field results for this native caterpillar, we used sentinel insects [Galleria mellonella L. (Lepidoptera: Pyralidae)] to identify particular locations where EPNs were present in the field. Platyprepia virginalis caterpillars were caged at these sites but again showed no evidence of susceptibility to EPNs. Platyprepia virginalis caterpillars reduce their exposure to EPNs by spending their time in and above the litter rather than contacting the soil when given the choice in nature. We conclude that P. virginalis is unlikely to serve as a reservoir for EPNs and that nematodes are unlikely to be important mortality factors for P. virginalis in this natural system.

Recently plant biologists have documented that plants, like animals, engage in many activities that can be considered as behaviors, although plant biologists currently lack a conceptual framework to understand these processes. Borrowing the well-established framework developed by psychologists, we propose that plant behaviors can be constructively modeled by identifying four distinct components: (1) a cue or stimulus that provides information, (2) a judgment whereby the plant perceives and processes this informative cue, (3) a decision whereby the plant chooses among several options based on their relative costs and benefits, and (4) action. Judgment for plants can be determined empirically by monitoring signaling associated with electrical, calcium, or hormonal fluxes. Decision-making can be evaluated empirically by monitoring gene expression or differential allocation of resources. We provide examples of the utility of this judgment and decision-making framework by considering cases in which plants either successfully or unsuccessfully induced resistance against attacking herbivores. Separating judgment from decision-making suggests new analytical paradigms (i.e., Bayesian methods for judgment and economic utility models for decision-making). Following this framework, we propose an experimental approach to plant behavior that explicitly manipulates the stimuli provided to plants, uses plants that vary in sensory abilities, and examines how environmental context affects plant responses. The concepts and approaches that follow from the judgment and decision-making framework can shape how we study and understand plant-herbivore interactions, biological invasions, plant responses to climate change, and the susceptibility of plants to evolutionary traps.

We examined genotypic variation in constitutive and induced resistance in grapes against Willamette spider mites, Eotetranychus willametti Ewing, and Pacific spider mites, Tetranychus pacificus McGregor, 2 common species of tetranychid mites found in California vineyards. We found evidence that early-season injury by Pacific mites induced resistance against subsequent Willamette mite populations but early-season injury by Willamette mites did not induce resistance against subsequent Willamette mite populations. Significant levels of induction were detected for several cultivars of the Old World species Vitis vinifera L. as well as the North American species V. calif arnica Bentham. Phylogenetic relationships among grape genotypes explained little of the variation we observed in induced resistance. Phylogenetic relatedness among grapes did help explain patterns of constitutive resistance for Pacific mites; cultivars of V. vinifera L. tended to be susceptible, whereas North American species were resistant. Wi11amette mites, however, performed well on some Old World cultivars and 2 North American species of Vitis that are native to California. We did not find any strong evidence of a negative correlation between constitutive resistance and strength of induction for these grape genotypes. Our results show that several factors contribute to variation in constitutive and induced resistance in grapes against these 2 species of spider mites, including grape genotype, previous history of mite injury (induction), the species of mite causing previous injury, and to some extent, phylogenetic relatedness among grapes. We also suspect that mite genotype has important influences.

Anti-herbivore defense shows high levels of both inter- and intraspecific variability. Defending against herbivores may be costly to the plant when it requires a tradeoff in allocation between defense and other missed opportunities, such as reproduction. Indeed, the plastic expression of defensive traits allows the plant to invest resources in defense only when the risk of being damaged actually increases, avoiding wasted resources. Plants may assess risk by responding to volatile cues emitted by neighbors that are under attack. Most plastic responses likely depend on environmental conditions. In this experiment, we investigated the effect of water availability on resistance induced by volatile cues in sagebrush. We found that plants receiving additional water over summer and/or volatile cues from neighbor donor plants showed reduced herbivore damage compared to control plants. Interestingly, we found no evidence of interactions between additional water and volatile cues. We performed an inferential analysis comparing historical records of the levels of herbivore damage during different years that had different temperature and precipitation accumulations. Results confirmed findings from the experiment, as the regression model indicated that sagebrush was better defended during wetter and hotter seasons. Reports from the literature indicated that sagebrush is extremely sensitive to water availability in the soil. We suggest that water availability may directly affect resistance of herbivory as well as sensitivity to cues of damage. Costs and benefits of allocating resources to defensive traits may vary with environmental conditions.

The possibility of communication between plants was proposed nearly 20 years ago, although previous demonstrations have suffered from methodological problems and have not been widely accepted. Here we report the first rigorous, experimental evidence demonstrating that undamaged plants respond to cues released by neighbors to induce higher levels of resistance against herbivores in nature. Sagebrush plants that were clipped in the field released a pulse of an epimer of methyl jasmonate that has been shown to be a volatile signal capable of inducing resistance in wild tobacco. Wild tobacco plants with clipped sagebrush neighbors had increased levels of the putative defensive oxidative enzyme, polyphenol oxidase, relative to control tobacco plants with unclipped sagebrush neighbors. Tobacco plants near clipped sagebrush experienced greatly reduced levels of leaf damage by grasshoppers and cutworms during three field seasons compared to unclipped controls. This result was not caused by an altered light regime experienced by tobacco near clipped neighbors. Barriers to soil contact between tobacco and sagebrush did not reduce the difference in leaf damage although barriers that blocked air contact negated the effect.

Trichomes are an important physical resistance mechanism of plants, as they reduce insect herbivore movement, feeding, and digestion. However, we know little about how trichomes influence herbivore distributions and populations. We conducted laboratory and field experiments to evaluate the preferences of Platyprepia virginalis (Boisduval) (Lepidoptera: Arctiidae) caterpillars to natural and manipulated densities of trichomes on their primary food, Lupinus arboreus Sims (Fabaceae). We then conducted field surveys to determine whether variation in trichome density among lupine bushes affected caterpillar spatial distribution on the landscape. Platyprepia virginalis caterpillars preferred lupine leaves with fewer trichomes in choice and no-choice experiments. In the field, caterpillar feeding damage was found more often on leaves with fewer trichomes. These preferences scaled up to the level of bushes in the landscape such that more caterpillars were found on bushes with lower trichome densities than on bushes with higher trichome densities. This is one of few studies to show the potential for trichome density to influence herbivore population size and distribution in a natural system at a landscape level. The results are consistent with trichomes functioning as a resistance mechanism with consequences for herbivore choice, performance, and distribution.

Mountain beavers cache plants in neat piles above the ground near the entrances to their underground burrows, although the purpose of these hay piles remains controversial. We tested the hypothesis that drying plants in piles above ground may slow decay when the plants are stored underground. Predried plants and undried controls were placed in abandoned burrows. Plants that had been predried retained more green and decayed more slowly than undried controls. This experimental test supports the hypothesis that haying may extend the time during which plants are useful to mountain beavers.

Summary There is substantial evidence that the rate of litter decomposition is affected by the match between the litter substrate and the soil matrix (decomposer community). We introduce and test the phenology–substrate-match hypothesis, which predicts that both litter composition and soil matrix will change over the course of the year and that a lagged match between litter type and soil matrix will result in an optimal decomposition environment. We conducted a decomposition experiment in a Mediterranean mixed deciduous–evergreen oak savanna in California. We initiated litter decomposition of both a deciduous oak (whose leaves fall in autumn) and an evergreen oak (whose leaves fall in spring) in both autumn and spring. Consistent with the phenology–substrate-match hypothesis, we found that decomposition of deciduous oak litter was accelerated compared to evergreen oak litter when decomposition was initiated in spring, while evergreen litter was accelerated compared to deciduous litter when decomposition was initiated in autumn. We also found a small effect of microsite on leaf decomposition, where both evergreen and deciduous oak leaves decomposed faster under the canopy of a conspecific. Synthesis. Our study extends theory of litter quality and the decomposer community into a temporal context, which may be an important source of variation in decomposition rates when species with different litterfall phenologies co-occur.

It is becoming increasingly accepted that plant–plant signals can affect plant defense against herbivores. In a few taxa, both the mechanisms of plant–plant signaling and the ecological consequences are well described. However, the current adaptive explanations for plant–plant signaling rely heavily on many ecological contingencies, such as the behavior of herbivores and genetic structure of plant neighborhoods, making it unclear how frequently plant–plant signaling is important across the plant kingdom. Moreover, these same ecological contingencies mean that it may be beneficial for a plant to release a publicly available cue to coordinate its (or its neighbor's) defenses following herbivory in some cases, but it may be disadvantageous in others. In this paper, we focus on recent studies that expand the taxonomic breadth of plant–plant signaling about herbivores that bring classic plant–plant signaling systems into a broader ecological context, and those that find either no consequence or counte...

As human-aided range expansions and climate change alter the distributions of plants and their herbivores, predicting and addressing novel species interactions will become increasingly pressing for community ecologists. In this context, a key, surprisingly understudied question is: when an exotic plant is introduced, which herbivores will adopt this new potential host? Whether the plant is a weed, an ornamental, or a crop, the development versus non-development of a novel plant – insect interaction can have profound eff ects for both economic and conservation applications. In this paper, we sketch mechanistic and statistical frameworks for predicting these interactions, based on how plant and herbivore traits as well as shared evolutionary history can infl uence detection, recognition, and digestion of novel plants. By emphasizing mechanisms at each of these steps, we hope to clarify diff erent aspects of novel interactions and why they may or may not occur. We also emphasize prediction and forecasting, as a major goal is to know in advance which interactions will develop from the many plant or insect introductions that occur in natural and man-made systems.

The tritrophic model featuring plants consumed by herbivores consumed by parasitoids or predators has become the primary paradigm used to describe herbivore dynamics. However, interactions involving herbivores can be habitat- specific and plants often provide habitat, as well as food. Structural complexity of the habitat may favor predators or may allow herbivore prey to escape detection and capture. This study considered the spatial and temporal dynamics of an arctiid caterpillar, Platyprepia virginalis. The tritrophic model that includes only a tachinid parasitoid that attacks P. virginalis and the caterpillars’ primary host-plant, Lupinus arboreus, has failed to provide much insight into this system. Instead, we found that ants killed and removed many small caterpillars. Protecting caterpillars from ants increased their survival three-fold and five-fold in assays conducted during two years. Caterpillars were more likely to survive in short-term assays at sites that naturally had a deeper cover of dead and living plant material. Experiments with baits showed that ant recruitment declined as litter depth increased on average. These survey results indicated that ant predation was an important source of mortality for young caterpillars and that the presence of thick litter reduced this mortality. These results were corroborated in an experiment that manipulated litter depth and ant access to caterpillars. Previous findings that other defoliating caterpillars increased litter depth and benefitted P. virginalis are also consistent with this hypothesis. Litter acts as an important non-trophic resource, allowing caterpillars to avoid predation by ants such that wet sites with deep litter act as source populations for caterpillars. Our results show strong effects of both trophic and non-trophic interactions since plants indirectly provided limiting habitat and this heterogeneous habitat strongly affected risk of predation and ultimately caterpillar abundance and distribution.

© 2016 by the Ecological Society of America. Plants respond to volatile cues emitted by damaged neighbors to increase their defenses against herbivores. We examined whether plants communicated more effectively with local neighbors than distant neighbors in a reciprocal experiment at two sites. Three branches on focal plants were incubated with air from (1) a control, (2) an experimentally clipped foreign plant from 230 km away, or (3) an experimentally clipped local plant from the same population as the focal plant. Branches incubated with air from the controls experienced 50-80%25 more leaf damage than those receiving air from experimentally clipped plants. Of more interest, branches receiving volatiles from experimentally clipped local plants received 50-65%25 of the leaf damage as those receiving volatiles from experimentally clipped foreign plants. Sabinyl compounds and related terpinenes were found to differ consistently for plants from southern and northern sites. These results indicate that cues vary geographically in their effectiveness and suggest that sagebrush responds more strongly to local than foreign dialects.

Plants that are damaged by herbivores emit complex blends of volatile compounds that often cause neighboring branches to induce resistance. Experimentally clipped sagebrush foliage emits volatiles that neighboring individuals recognize and respond to. These volatiles vary among individuals within a population. Two distinct types are most common with either thujone or camphor as the predominate compound, along with other less common types. Individuals respond more effectively to cues from the same type, suggesting that some of the informative message is contained in the compounds that differentiate the types. In this study, we characterized the chemical profiles of the two common types, and we examined differences in their microhabitats, morphologies, and incidence of attack by herbivores and pathogens. Analysis by gas chromatography coupled with mass spectrometry revealed that the camphor type had higher emissions of camphor, camphene, and tricyclene, while the thujone type emitted more α-thujone, β-thujone, (Z)-salvene, (E)-salvene, carvacrol, and various derivatives of sabinene. We were unable to detect any consistent morphological or microhabitat differences associated with the common types. However, plants of the thujone type had consistently higher rates of damage by chewing herbivores. One galling midge species was more common on thujone plants, while a second midge species was more likely to gall plants of the camphor type. The diversity of preferences of attackers may help to maintain the variation in volatile profiles. These chemical compounds that differentiate the types are likely to be informative cues and deserve further attention.

Summary 1. There are now approximately 10 examples of plants that use volatile cues emitted by damaged neighbours to adjust their defences against herbivores. For two of these examples, preliminary evidence suggested that plants may experience net benefits from such eavesdropping, although eavesdropping was uncommon in one case and estimates of plant fitness were ambiguous in the other case. 2. In the current study, we examined the long-term consequences of exposure to cues emitted by experimentally clipped sagebrush neighbours. In this sagebrush system we have repeatedly found that sagebrush plants that have experimentally clipped neighbours experience less herbivore damage over the season than plants with unclipped control neighbours under field conditions. We followed a cohort of young sagebrush plants from emergence in 1999 for 12 years. Neighbours of half of these plants were artificially clipped every spring from 2004–08 and survival and flowering was measured in each autumn from 1999–2011. 3. Survival of marked branches of young plants was not consistently affected by whether its neighbour was clipped. Plants near clipped neighbours produced more branches during this period than those near unclipped neighbours. There were no measurable treatment effects on plant survival over the 12 years. Branches near clipped neighbours produced more inflorescences than branches near unclipped neighbours. 4. Seedlings were more likely to survive to the end of their first dry season in two different years near clipped neighbours compared to unclipped neighbours. 5. Synthesis. The results suggest different effects of clipped neighbours that depend on plant age. Responding to the cues of experimental clipping may provide a slight net benefit, considering these results and other published studies, even though these cues provided little predictive value about actual risk of herbivory. Responding to reliable cues may be even more beneficial and may favour plants that eavesdrop on neighbours.

1. Ecologists have argued about the commonness and strength of interspecific competition between insect herbivores, but facilitation between herbivores has received much less consideration. We previously found that when two species of folivorous caterpillars co-occurred on a shared host plant, feeding by early season tiger moth caterpillars reduced the growth and reproduction of later season tussock caterpillars. However, densities of tussock caterpillars in summer were positively correlated with densities of tiger moth caterpillars the following spring. 2. In this study, we experimentally manipulated numbers of feeding tussock caterpillars and found that they facilitated tiger moth caterpillars. 3. The depth of the litter layer beneath host lupine bushes was positively correlated with the number of tussock caterpillars feeding on each bush. Experimental additions of litter beneath lupine canopies during summer resulted in increased numbers of tiger moth caterpillars in the following spring, indicating a causal role of litter. Litter potentially provides food, habitat and protection from desiccation and predation. We failed to find evidence that tussock caterpillars facilitated tiger moth caterpillars by mechanisms independent of litter. 4. Our study demonstrates that facilitation may operate between insect herbivores, across life-stages through indirect interactions that are non-trophic. Facilitation operated by a novel mechanism, the accumulation of litter which was a by-product of feeding by one species was valuable to a second species. Facilitation persisted in time and space far beyond the creation of litter by tussock caterpillars which should be considered important ecosystem engineers from the point of view of tiger moths. Facilitations that involve habitat modification may generally connect species that do not interact directly or trophically, and have not previously been considered to affect one another.

Volatile communication allows plants to coordinate systemic induced resistance against herbivores. The mechanisms responsible and nature of the cues remain poorly understood. It is unknown how plants distinguish between reliable cues and misinformation. Previous experiments in which clipped sagebrush branches were bagged suggested that cues are emitted or remain active for up to 3 days. We conducted experiments using plastic bags to block emission of cues at various times following experimental clipping. We also collected headspace volatiles from clipped and unclipped branches for 1 h, transferred those volatiles to assay branches, and incubated the assays for either 1 or 6 h. We found that assay branches that received volatile cues for less than 1 h following clipping of neighbors failed to induce resistance. Assay branches that received volatile cues for more than 1 h experienced reduced herbivory throughout the season. Branches incubated for 6 h with volatiles that had been collected during the first hour following clipping showed induced resistance. These results indicate that sagebrush must receive cues for an extended time (>1 h) before responding; they suggest that the duration of cue reception is an important and overlooked process in communication allowing plants to avoid unreliable, ephemeral cues.

Plants can release chemicals into the environment that suppress the growth and establishment of other plants in their vicinity: a process known as 'allelopathy'. However, chemicals with allelopathic functions have other ecological roles, such as plant defense, nutrient chelation, and regulation of soil biota in ways that affect decomposition and soil fertility. These ecosystem-scale roles of allelopathic chemicals can augment, attenuate or modify their community-scale functions. In this review we explore allelopathy in the context of ecosystem properties, and through its role in exotic invasions consider how evolution might affect the intensity and importance of allelopathic interactions.

Legacy effects occur when particular species or their interactions with others have long-lasting impacts, and they are increasingly recognized as important determinants of ecological processes. However, when such legacy effects have been explicitly explored, they most often involve the long-term direct effects of species on systems, as opposed to the indirect effects. Here, we explore how a legacy of small mammal exclusion on the abundance of a shrub, bush lupine (Lupinus arboreus), influences the abundance of a native land snail (Helminthoglypta arrosa) in coastal prairie and dune habitats in central California. The factors that limit populations of land snails are very poorly known despite the threats to the persistence of this group of species. In grasslands, prior vole (Microtus californicus) exclusion created long-lasting gains in bush lupine abundance, mediated through the seedbank, and was associated with increased snail numbers (10×) compared to control plots where mammals were never excluded. Similar plots in dune habitat showed no difference in snail numbers due to previous mammal exclusion. We tested whether increased competition for food, increased predation, and/or lower desiccation explained the decline in snail numbers in plots with reduced lupine cover. Tethering experiments supported the hypothesis that voles can have long-lasting impacts as ecosystem engineers, reducing woody lupine habitat required for successful aestivation by snails. These results add to a growing list of studies that have found that non-trophic interactions can be limiting to invertebrate consumers.