Natural grasslands are one of the most threatened ecosystems in North America. Conservation efforts are often hampered by exotic plant invasions in existing remnant natural grasslands. Tall fescue [Schedonorus phoenix (Scop.) Holub.] is an introduced cool-season (C3) forage/turf grass which readily invades natural grasslands in Kentucky and neighboring states. Our study objectives were to (1) compare the efficacy and application timing effect of clethodim to that of imazapic to selectively remove tall fescue from natural grasslands and (2) evaluate the response of the nontarget grassland plant community (i.e., native grass and forb canopy cover) following herbicide treatments. Clethodim and imazapic treatments consisted of early (April 4, 2001) and late (April 20, 2001) applications, and these were applied at 0.23 and 0.21 kg ai ha−1, respectively. Both herbicides reduced tall fescue cover (P ≤ 0.05); herbicide application timing had no effect on herbicide efficacy to control tall fescue. Native grass cover was higher (P ≤ 0.05) in all herbicide-treated plots compared to the untreated controls, except for the late clethodim-treated plots. All herbicide treatments increased forb abundance compared to controls. Spring applications of clethodim were equally effective to those of imazapic at controlling tall fescue in natural grasslands. Imazapic released native grasses better than clethodim, whereas clethodim was better at increasing forb abundance. Furthermore, early clethodim treatments had fewer nontarget effects on native C4 grasses compared to late clethodim treatments. Overall, clethodim shows promise as a beneficial management tool for tall fescue control in C4-dominated natural grasslands.

The invasive aquatic plant hydrilla rapidly spread through the 28,500 ha Kissimmee Chain of Lakes (KCOL) system in Florida in the late 1980s and early 1990s. Large-scale herbicide treatments with fluridone were initiated in 1993 and resulted in widespread reduction in hydrilla; however, by 2000, sustained use of fluridone resulted in dominance of fluridone-resistant strains of hydrilla throughout these lakes. The last large-scale fluridone applications on the KCOL were conducted in 2004, and in 2012, a sampling effort was initiated to determine the status of fluridone-resistant strains of hydrilla given an 8-yr period with no further selection pressure from fluridone. A total of 260 sites were sampled on the lakes during March, May, September, and December 2012. Plants were returned to the lab and exposed to fluridone at concentrations of 5, 10, and 20 μg L−1 and a pulse-amplitude-modulated (PAM) fluorometer was utilized to measure fluorescence yield of new shoot tissue growth following a 14-d exposure period. Results indicate that 80 to 90% of the sites sampled on the four lakes of the Kissimmee Chain remain resistant to fluridone. Three distinct patterns of response to fluridone were noted, suggesting that susceptible, moderately tolerant, and highly tolerant strains of hydrilla currently coexist on these lakes. Although fluridone-susceptible plants were present on the KCOL, this study clearly demonstrates that most of the hydrilla remained resistant despite an 8-yr period with no fluridone selection pressure.

Biological control is one of the most common approaches used to manage invasive weeds of wetlands and other natural areas. Before candidate agents can be released, research is conducted to support biological control, which can be protracted and expensive, leading to a scientific and potentially lengthy regulatory review. To increase biological control safety, efficacy, and transparency, we suggest that during the early phases of a weed project, the feasibility of the invasive plant as a target should be studied explicitly. Our purpose here is to summarize information of an important invasive weed that can serve to judge whether the project is appropriate. Chinese tallowtree, Triadica sebifera, is one of the worst invasive species invading coastal wetlands and other riparian areas of the southeastern United States. Current management practices have not controlled the spread of this weed into these sensitive habitats. Initial surveys in the plant's native Chinese range for potential biological control agents have recovered several herbivore species that could be developed. These potential agents include defoliators, root and foliage feeders, and gall formers, whose biology, apparent host specificity, and impacts on plant fitness suggest that biological control offers great promise against Chinese tallowtree. When conducted during the initial phase of a project, this type of feasibility study can address potential conflicts of interest and risks, ultimately producing projects that are more effective and safer for biological control.

Corallita (Antigonon leptopus) is a perennial vine, lauded as an ornamental for its vigorous growth, and plentiful (usually) pink flowers, and even its ability to smother unsightly landscapes. In the United States it thrives in horticultural zones 8 to 10, and also is successfully grown worldwide in tropical climates. When corallita is neglected, it can grow quickly over other vegetation, spreading beyond its area of introduction. Once established, it is difficult to eradicate because it produces many tuberous roots that can propagate vegetatively. Its fruits are buoyant, allowing for successful seed dispersal in water. The islands of Guam (South Pacific Ocean) and St. Eustatius (Caribbean Sea) represent two regions where corallita has become so pervasive that it threatens local diversity. In Florida, already it is classified as a Category II invasive. Our report reviews the literature and past studies of corallita, in addition to adding new taxonomic and distribution information from herbarium specimens to clarify the identity and geographic range. It is recommended that introductions of this plant by the horticultural industry in both tropical and temperate regions be closely monitored to prevent spread. On tropical island nations, we advise against any new introductions.

Diversified grasslands that contain native plant species can produce biofuels, support sustainable grazing systems, and produce other ecosystem services. However, ecosystem service production can be disrupted by invasion of exotic perennial plants, and these plants can have soil-microbial “legacies” that may interfere with establishment and maintenance of diversified grasslands even after effective management of the invasive species. The nature of such legacies is not well understood, but may involve suppression of mutualisms between native species and soil microbes. In this study, we tested the hypotheses that legacy effects of invasive species change colonization rates, diversity, and composition of arbuscular-mycorrhizal fungi (AMF) associated with seedlings of co-occurring invasive and native grassland species. In a glasshouse, experimental soils were conditioned by cultivating three invasive grassland perennials, three native grassland perennials, and a native perennial mixture. Each was grown separately through three cycles of growth, after which we used T-RFLP analysis to characterize AMF associations of seedlings of six native perennial and six invasive perennial species grown in these soils. Legacy effects of soil conditioning by invasive species did not affect AMF richness in seedling roots, but did affect AMF colonization rates and the taxonomic composition of mycorrhizal associations in seedling roots. Moreover, native species were more heavily colonized by AMF and roots of native species had greater AMF richness (number of AMF operational taxonomic units per seedling) than did invasive species. The invasive species used to condition soil in this experiment have been shown to have legacy effects on biomass of native seedlings, reducing their growth in this and a previous similar experiment. Therefore, our results suggest that successful plant invaders can have legacies that affect soil-microbial associations of native plants and that these effects can inhibit growth of native plant species in invaded communities.

Fire is unlikely to be a simple, deterministic process that will result in increased rates of invasion by nonnative plants. Relative dominance of nonnative plants at any postfire site likely results from a combination of factors, including propagule pressure (aboveground and belowground), time since invasion, interspecific competition, disturbance history, rainfall patterns, soil nutrients, environmental heterogeneity, land use, plus the actual fire dynamics. Consequently, the role different factors play in nonnative plant invasion is still relatively unexplained, and the possibility that wildfire may promote nonnative plant species remains a concern. Our aim was to evaluate the state of knowledge concerning how nonnative plant species establish, survive, and spread following wildfire in wildland areas for the main vegetation types of the Intermountain West. Although the particular invading species varied with vegetation type, the most frequently reported species were annual grasses (e.g., downy brome [Bromus tectorum], Japanese brome [Bromus japonicus]) and long-distance, dispersing forbs (e.g., Canada thistle [Cirsium arvense], common dandelion [Taraxacum officinale], and prickly lettuce [Lactuca serriola]). Overall, many of the reviewed studies found an increase, followed by a long-term decline, in nonnative species after wildfire, particularly when other disturbances were minimal. Regardless of this finding, many people consider rapid detection and control of new, nonnative plant populations after wildfire one of the most effective management activities. That axiom may be correct, if the nonnative propagule pressure is high and the anthropogenic disturbances are frequent, but if this is not the case the question to answer is whether immediate management, including reseeding, will benefit the system or whether off-target effects could make that approach undesirable. Continued investigation and increased quantitative analysis of the complex factors that likely contribute to changes in nonnative plants dynamics following wildfire should improve our understanding of the issue and assist managers in allocating limited resources.

The invasive grass Bromus tectorum (cheatgrass, downy brome) has extensive ecological breadth in its introduced range, and is increasing in abundance at high elevations. As a predominantly selfing species with high potential population growth rates, it is a likely candidate for having general-purpose genotypes that facilitate its invasion into high elevations. We evaluated evidence for general-purpose genotypes by examining phenotypic and molecular variation in B. tectorum from low- and high-elevation sites in the Rocky Mountains. Common garden experiments revealed that plants from high elevations germinated faster, but no other phenological, morphological, or physiological differences were found between elevations. Leaf length was longer for low-elevation sites in a common garden using field-collected seeds, but this was not found in a common garden using greenhouse-grown seeds, suggesting maternal effects. For most traits, there was significant phenotypic variation among sites and individuals. Using microsatellites, we genotyped individuals from low- and high-elevation sites. We found no indication of genetic differentiation at these loci between high and low elevations, but there was significant variation within and among sites. No single genotype was common across sites. The success of B. tectorum is not due to a few general-purpose genotypes, but rather many genotypes associated with many different phenotypes. Understanding the mechanisms behind the success of invaders enhances our knowledge of basic evolutionary and ecological processes and is crucial to our ability to manage ongoing, and prevent future, invasions, especially into new habitats.

Prescribed burning can control invasive annual grasses that threaten the biological and economic value of California grasslands. Susceptibility of grass seed to burning can depend on burn timing, exposure time, and type of exposure (direct flame heat or convective heat); thus, these factors can influence the success of a prescribed burning program. To further investigate these factors, laboratory simulations were conducted on barb goatgrass, medusahead, and ripgut brome at several stages of seed maturity, as determined by percent moisture of the inflorescences. Seeds were exposed either to direct flame using a Bunsen burner or to heated air in a muffle furnace. Flame treatments were conducted at one temperature (∼400 C) and several exposure times (0 to 14 s), depending on the species. Furnace treatments included four temperatures (150, 200, 250, and 300 C) and seven exposure times (0, 10, 20, 30, 40, 60, or 80 s). Seed germination was analyzed for each temperature series to determine the LD50 and LD90 in seconds of exposure time. Susceptibility to furnace treatments, which simulated heat exposure of seeds on the soil surface, was not statistically different within a range of seed moisture levels for all three species. The LD50 values at 250 C (typical soil temperature with grassland fire) ranged from 28 to 49 s, which far exceeds the time of exposure during a typical grassland fire. Susceptibility to flame showed a similar lack of change over maturation of medusahead and barb goatgrass seeds, with LD90 values ranging between 4.8 and 7.4 s for all seed moisture levels. In contrast, ripgut brome seeds exposed to flame showed increasing susceptibility with reduced seed moisture content. The LD90 values for exposure were less than one second for seed moisture levels at or below 10%, compared to 3.7 s for seeds at 55 to 60%. Although flame susceptibility increased for ripgut brome, seeds at all maturation stages were more sensitive than medusahead and barb goatgrass. Additionally, the LD90 values for all three species are attainable under field conditions. Thus, burn prescriptions for these three species are not constrained by maturation stage, but should occur prior to seed drop and when fuel loading is high. This will maximize exposure time of seeds to direct flame.

Effective control techniques for pale swallowwort (PSW), an invasive herbaceous vine of old fields and forest understories, are limited. We conducted a 3-yr cutting and herbicide study on an adjacent old-field and forest understory site near Ithaca, NY, for control of PSW. Plants in experimental plots were cut in early July and cut again or sprayed in late August for two seasons with the isopropylamine salt of glyphosate, or one of two rates (low or high) of either triclopyr triethylamine salt (i.e., SL, SH) or triclopyr butoxyethyl ester (EL, EH). The herbicide treatments were effective in reducing PSW cover, plant (stem) density, and aboveground biomass in the old-field site, but in several cases, only after 2 yr of cutting plus herbicide application. Only the cutting plus SH treatment did not reduce PSW cover relative to the unmanaged control in the forest understory and no treatment reduced biomass. In general, the cutting plus EH treatment was most effective in reducing PSW stem densities in the forest site. The most effective herbicide treatments differed between sites. Cutting plus EH reduced PSW cover by 84% and stem density (> 5 cm) by 86% in the old-field site. Cutting plus SH effectively decreased long and short (≤ 5 cm) stem densities by 86 and 96%, respectively. Cutting plants twice during each of two seasons increased PSW cover by 301% and density of stems > 5 cm by 73% at this site. In the forest site, cutting plus glyphosate, or cutting plus EH or cutting plus SL and EL resulted in the greatest reductions in PSW cover (80, 76, 66, and 56%, respectively). Cover in plots cut twice per year decreased by 19%. The EH or SL treatments decreased long-stem densities by 78 and 71%, respectively. The EH treatment decreased short-stem density by 37%. These findings suggest that integrated techniques may control PSW but that effective management strategies may be habitat constrained.

Prairies are imperiled habitats, with remnants being generally small and often existing in isolation. Invasive plants have the potential to invade not just the edge of small remnants but also the interior because smaller remnants experience greater edge effects than do large, contiguous prairies. Additionally, invasive plants limit recruitment of native plants, which can arrest secondary succession. We proposed to assess techniques for restoration that included removing annual grasses and supplementing native species recruitment with seeding of native grass and forb species. We also assessed the effect of specific factors affecting recruitment: soil moisture and seed predation. Treatments included broadcast, spot, or no application of the herbicides imazapic and glyphosate and with or without seeding plus mulch. With treatments nested within each of three plant communities, ranging from annual- to perennial-dominated communities, in four blocks per community, plant characteristics (percentage of cover and plant density), soil moisture availability, and seed-predation losses were measured along a plant community gradient within one season at two locations. A combination of broadcast herbicide application and seeding with mulching was found to be more effective in reducing annual grasses and enhancing the establishment of native grass species in predominately annual and mixed communities (annuals and perennials). Spot herbicide application was effective in predominately perennial communities, whereas only seeding native species did not improve recruitment. Although seed predation reduced seedling recruitment, mulch provided seed protection and enhanced soil moisture retention. Plant community response to imposed treatments differed among communities, suggesting that a decision support tool would facilitate management decisions tailored for each plant community. The decision tool would be useful to ensure that appropriate treatments are applied and that specific factors affecting recruitment, such as seed predation and soil moisture, are addressed.

Null hypothesis significance testing (NHST) forms the backbone of statistical inference in invasive plant science. Over 95% of research articles in Invasive Plant Science and Management report NHST results such as P-values or statistics closely related to P-values such as least significant differences. Unfortunately, NHST results are less informative than their ubiquity implies. P-values are hard to interpret and are regularly misinterpreted. Also, P-values do not provide estimates of the magnitudes and uncertainties of studied effects, and these effect size estimates are what invasive plant scientists care about most. In this paper, we reanalyze four datasets (two of our own and two of our colleagues; studies put forth as examples in this paper are used with permission of their authors) to illustrate limitations of NHST. The re-analyses are used to build a case for confidence intervals as preferable alternatives to P-values. Confidence intervals indicate effect sizes, and compared to P-values, confidence intervals provide more complete, intuitively appealing information on what data do/do not indicate.

The objective of this article is to identify growth patterns of Japanese knotweed propagules distributed by high-water events. Along four river systems, we collected and measured Japanese knotweed propagules that had been distributed by flooding approximately 1 yr earlier. Results indicate that the size of the emergent shoot may be determined by the extent of underground growth late in the growing season, although initially it is linked to the size of the propagule. Our results show that 70% of new plants originated from rhizome fragments, and 30% from stems. This proportion is similar to regeneration rates shown in laboratory studies. We suggest that the best way to prevent the spread of Japanese knotweed along rivers is to focus control efforts on those stands most susceptible to erosion and propagule dispersal. We also suggest that an early detection and rapid response management approach can be effectively utilized to eradicate these propagules, and effectively suppress the spread of Japanese knotweed. Our data-collection method also provides evidence that control of newly distributed propagules can be effectively accomplished without the use of herbicides or heavy mechanical tools.

The identification of privet in the middle southern United States can be difficult. Because most introduced species of privet can be invasive, and recent mapping projects seek location and species population data, proper identification is important. Without proper identification of privet species, data on species distributions and other pertinent information regarding invasiveness could lead to improper conclusions. Currently, information on privet identification is scattered throughout a number of reference materials. The purpose of this publication is to assist with the proper identification of escaped privet species, and suggest management options.

Invasive species with distributions that encompass much of the North American environment often demand a range of management approaches, for several key reasons. Firstly, the North American environment includes a large number of highly variable habitats in terms of climatic, edaphic, and landscape features. Secondly, these regional habitat differences are accentuated by jurisdictions within Canada and the United States, whereby approaches and available resources differ at local, regional, and national scales. Another important consideration is whether an invasive species or complex also possesses genetic variation. All three of these factors render the knotweed complex in North America a highly variable target for management. In this paper we review existing knowledge of the variable nature of knotweed species (Fallopia japonica (Houtt.) Ronse Decr., Fallopia sachalinensis (F. Schmidt ex Maxim) Ronse Decr., and Fallopia × bohemica, (Chrtek and Chrtková) J. P. Bailey in North America, and evaluate how herbicidal, mechanical and biological control measures must account for this genetic variation, as well as accounting for regional differences and the potential northward expansion of knotweed under climate change. The imminent release of the psyllid, Aphalara itadori Shinji as a biological control agent in North America must also navigate regional and genetic differences. Prior European experience dealing with the three knotweed species should prove useful, but additional research is needed to meet the emerging challenge posed by F. × bohemica in North America, including the possibility of glyphosate resistance. Managers also face challenges associated with posttreatment restoration measures. Furthermore, disparities in resources available to address knotweed management across the continent need to be addressed to contain the rapid spread of this highly persistent and adaptable species. Linking practitioners dealing with knotweed “on the ground” with academic research is a crucial step in the process of marshalling all available resources to reduce the rapidly spreading populations of knotweed.

Buffelgrass, a highly competitive and flammable African bunchgrass, is spreading rapidly across both urban and natural areas in the Sonoran Desert of southern and central Arizona. Damages include increased fire risk, losses in biodiversity, and diminished revenues and quality of life. Feasibility of sustained and successful mitigation will depend heavily on rates of spread, treatment capacity, and cost–benefit analysis. We created a decision support model for the wildland–urban interface north of Tucson, AZ, using a spatial state-and-transition simulation modeling framework, the Tool for Exploratory Landscape Scenario Analyses. We addressed the issues of undetected invasions, identifying potentially suitable habitat and calibrating spread rates, while answering questions about how to allocate resources among inventory, treatment, and maintenance. Inputs to the model include a state-and-transition simulation model to describe the succession and control of buffelgrass, a habitat suitability model, management planning zones, spread vectors, estimated dispersal kernels for buffelgrass, and maps of current distribution. Our spatial simulations showed that without treatment, buffelgrass infestations that started with as little as 80 ha (198 ac) could grow to more than 6,000 ha by the year 2060. In contrast, applying unlimited management resources could limit 2060 infestation levels to approximately 50 ha. The application of sufficient resources toward inventory is important because undetected patches of buffelgrass will tend to grow exponentially. In our simulations, areas affected by buffelgrass may increase substantially over the next 50 yr, but a large, upfront investment in buffelgrass control could reduce the infested area and overall management costs.

Miscanthus sinensis is a perennial grass native to Asia, but since its introduction to the United States in the late 19th century, it has become both a major ornamental crop and invasive species. Previous studies of the ecology of M. sinensis in both its introduced and native ranges have suggested that it may be occupying a novel ecological niche in the introduced range. Miscanthus sinensis and its daughter species, Miscanthus × giganteus, are under evaluation as bioenergy crops; therefore, characterization of the ecology and environmental niche of M. sinensis is essential to mitigate the risk of fostering future invasion in the United States. In July 2011, we surveyed 18 naturalized M. sinensis populations spanning the U.S. distribution, covering a 6° latitudinal gradient from North Carolina to Massachusetts. Miscanthus sinensis populations ranged in size from 3 to 181,763 m2 with densities between 0.0012 and 2.2 individuals m−2, and strongly favored highly disturbed and unmanaged habitats such as roadsides and forest edges. Population size and individual plant morphology (i.e., tiller height, basal diameter, and tiller number) were not affected by soil characteristics and nutrient availability, though increased tree canopy cover was associated with reduced population size (P < 0.0001). Plant size and vigor were not significantly affected by low light availability, which supports previous suggestions of shade tolerance of M. sinensis. In summary, M. sinensis can tolerate a broad range of climatic conditions, light availability, and nutrient availability in the eastern United States, suggesting risk of further invasion beyond its current distribution in the United States.

Tamarisk species are shrubs or small trees considered by some to be among the most aggressively invasive and potentially detrimental exotic plants in the United States. Although extensively studied in the southern and interior west, northwestern (Oregon, Washington, and Idaho) distribution and habitat information for tamarisk is either limited or lacking. We obtained distribution data for the northwest, developed a habitat suitability map, and projected changes in habitat due to climate change in a smaller case study area using downscaled climate data. Results show extensive populations of tamarisk east of the Cascade Mountains. Despite the perceived novelty of tamarisk in the region, naturalized populations were present by the 1920s. Major population centers are limited to the warmest and driest environments in the central Snake River Plain, Columbia Plateau, and Northern Basin and Range. Habitat suitability model results indicate that 21% of the region supports suitable tamarisk habitat. Less than 1% of these areas are occupied by tamarisk; the remainder is highly vulnerable to invasion. Although considerable uncertainty exists regarding future climate change, we project a 2- to 10-fold increase in highly suitable tamarisk habitat by the end of the century. Our habitat suitability maps can be used in “what if” exercises as part of planning, detection, restoration, management, and eradication purposes.

Kudzu is an invasive perennial climbing vine characterized by fast growth rates and tolerance to control measures. Repeated applications with high rates of 2,4-D plus picloram provide effective kudzu control, but picloram use is not permitted in certain states due to groundwater pollution concerns. Studies were conducted in Alabama and Florida to compare kudzu control with aminocyclopyrachlor, a new herbicide, to control provided by aminopyralid, clopyralid, metsulfuron methyl, and picloram plus 2,4-D, which are common treatments for kudzu management. Two annual applications of the same herbicide treatment were evaluated for effects on kudzu cover, kudzu volume index, and cover of other vegetation. Aminocyclopyrachlor at 140 to 280 g ae ha−1 (2 to 4 oz ae ac−1) was as effective as the standard 4.48 kg ae ha−1 (4 lb ae ac−1) 2,4-D amine plus 1.2 kg ae ha−1 picloram for kudzu control. There were no differences in kudzu control among the three rates of aminocyclopyrachlor tested. Colonization by graminoids, forbs, and Rubus spp. at 2 yr was greatest for herbicides providing the best kudzu control: aminocyclopyrachlor, and 2,4-D plus picloram. Herbicide treatments were more effective in controlling kudzu at the Alabama location, but repeated annual applications for 2 yr did not completely eliminate kudzu with any treatment at either site.

Chemical, mechanical, and biological methods are used to manage invasive plants, but their effectiveness at removing specific plant invaders while preserving native communities varies widely. Chemical methods are used most extensively but the nontarget effects of some herbicides can have lasting effects on native plants. Nonchemical methods are needed for sites containing rare or threatened native species and where the cost of herbicides is prohibitive. Here we evaluate multiple nonchemical methods for removing Japanese stiltgrass, a nonnative annual grass that is rapidly invading eastern U.S. forests. We applied mowing, hand weeding, and spring and fall fire treatments to replicated plots at three forested sites in southern Indiana and compared the response of Japanese stiltgrass and native plants to untreated reference plots. Mowing and fall fires applied just before seed set were the most effective methods for removing Japanese stiltgrass. Mowing decreased invader cover by 70% and biomass by 95%, whereas fall fires reduced cover by 79% and biomass by 90% compared to reference plots. Spring fire reduced Japanese stiltgrass cover, but not biomass, and hand weeding did not significantly reduce invader cover or biomass compared to untreated plots. There were no significant differences in the response of the overall native plant community or of specific native plant functional groups to the removal treatments. In summary, mowing and properly timed fall fires may be effective nonchemical methods for managing Japanese stiltgrass invasions and restoring native communities. Future research should focus on evaluating the responses of Japanese stiltgrass, native species, and other plant invaders to removal treatments conducted over successive growing seasons across a range of invaded habitats.

The effect of invasive species might be lessened if herbivores reduced transpiration and growth rates; however, simply removing photosynthetic material might not ensure that the transpiration rate of active leaf tissue decreases. We assessed whether biological control has an injurious effect on the target plant species, giant reed (Arundo donax), by quantifying leaf photosynthetic and transpiration responses to two herbivores: an armored scale, Rhizaspidiotus donacis, and a stem-galling wasp, Tetramesa romana. Herbivory by a sap-feeding scale and a stem-galling wasp both separately and together, reduces the rates of leaf level physiological processes in A. donax. The effect of the wasp increases with density and reduces photosynthesis by reducing the carboxylation rate of ribulose-1,5-bisphosphate carboxylase oxygenase, which controls CO2 fixation in photosynthesis. The scale insect reduces photosynthesis by decreasing the maximum rate of electron transport, which determines how much light energy can be captured in photosynthesis. The effect of the armored scale takes approximately 5 mo after infestation, which coincides with generation time. When both insects are present at the same time, the effect of their herbivory appears additive after time for the scale to reproduce. We conclude that a combination of two herbivores can have a stronger physiological effect than one type of herbivore, likely because of their different effects on leaf function.