Earthwatch 2015 Annual Field Report. CLIMATE CHANGE AND CATERPILLARS TROPHIC INTERACTIONS, BIODIVERSITY, AND CLIMATE CHANGE Lee Dyer, PhD

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1 Earthwatch 2015 Annual Field Report CLIMATE CHANGE AND CATERPILLARS TROPHIC INTERACTIONS, BIODIVERSITY, AND CLIMATE CHANGE Lee Dyer, PhD REPORT COMPLETED BY: LEE DYER, ANGELA SMILANICH, CHRIS JEFFREY, LORA RICHARDS, JANE DELL, ANDREA GLASSMIRE, NICK PARDIKES, DANIELLE SALCIDO, TOM WALLA ECOLOGY, EVOLUTION, AND CONSERVATION BIOLOGY, UNIVERSITY OF NEVADA, RENO EARTHWATCH EXPEDITIONS, 1-JUNE TO 31-DECEMBER,

2 Lee Dyer Professor and Director PhD program Ecology, Evolution and Conservation Biology 24-May, 2016 Dear Volunteers, Thanks to your hard work and interest, the 2015 field season was successful by any measure. It included multiple trips to 4 of our research sites and 6 excellent research teams from Ecuador up to the Great Basin. As a group, our teams continue documented complex relationships between climate, plants, plant chemistry, caterpillars, parasitoids, interaction diversity, and ecosystem services across deserts, forests, and grasslands in the Americas. I am extremely grateful for your help towards documenting biodiversity and discovering new network links, new phytochemicals, and new species of insects at all of our sites. One great accomplishment for 2015 was that we published 2 papers in the Proceedings of the National Academy of Sciences. Both papers utilized 18 years of Earthwatch data on caterpillars, host plants, and parasitoids to address these sorts of questions: What are the consequences of plants having high diversity of defensive toxins and how is this potentially affected by climate? How do caterpillars and parasitoids respond? For a special group of plants at the Costa Rica and Ecuador sites (the wild peppers) we found that the concentration and types of different toxins help maintain high caterpillar and parasitoid diversity and high levels of caterpillar and parasitoid specialization. The strong effects of climate on these toxins are likely to lead to outbreaks of damaging insects and contribute to lower levels of biodiversity. I hope that you are all able to find interesting information in this report, and hopefully it will provide you with a sense of how successful the Climate Change and Caterpillars project has been because of your hard work. I will always be impressed with the amount of high quality data that Earthwatch volunteers have collected with us over the past decades. In the face of continual bad news about increases in extreme weather events, rising sea levels, massive extinctions, and loss of our favorite habitats, it is important to continue the hard work of documenting and protecting the amazing diversity that lives on through the consequences of human impacts. My collaborators and I have enjoyed working with all of you, and our project would not be successful without your help. Please feel free to keep in touch and we will do our best to respond. Hopefully we'll see you again in the field. Sincerely, Lee Dyer, Professor, ; ldyer@unr.edu Ecology, Evolution, Conservation Biology University of Nevada, Reno/0314 Reno, Nevada

3 SUMMARY As a team of 7 Earthwatch expeditions, we collected over 1,000 individual caterpillars at 4 different sites, reared out over 200 individual parasitic flies and wasps, discovered new phytochemicals, discovered dozens of novel interactions, and published a dozen papers using years of accumulated Earthwatch data. Our papers demonstrate the complex mechanisms by which climate change can cause damaging decreases in biodiversity and again provide an impetus to document the vast unknown biodiversity on earth in this case plant chemical diversity and insect diversity. We made progress towards documenting this biodiversity and continued to discover new interactions, new phytochemicals, and new species of insects at all of our sites. GOALS, OBJECTIVES, AND RESULTS Objective 1: Determine how climate change parameters affect biodiversity. We continue to make progress towards our long term goals of determining effects of hurricanes and climatic variation on interaction diversity through our modeling approach and through empirical data (Scherrer et al. 2016). Our models, including analytical, simulation, and statistical approaches are being parameterized using all of the Earthwatch data from all of our sites (e.g., see Dyer and Forister 2016). One particularly exciting topic that is important for enhancing these models addresses questions that are seemingly unrelated to climate-change biodiversity loss: What are the consequences of plants having high diversity of defensive toxins and how is this potentially affected by climate? How do caterpillars and parasitoids respond? In 2015, we used 18 years of Earthwatch data to address these questions and published a paper in the Proceedings of the National Academy of Sciences about our results (Richards et al. 2015). For a special group of rainforest plants, the wild peppers (Piper spp., Piperaceae), we found that the concentration and types of different toxins help maintain high caterpillar and parasitoid diversity and high levels of caterpillar and parasitoid specialization. Previously, we had published results showing that potent toxins increase in response to higher temperatures and increasing CO 2 concentrations or lower phytochemical diversity (Dyer et al. 2013). Thus, it is likely that the high correlation between plant chemical diversity and insect diversity is being strongly disrupted by climate change parameters. Taken together, these published results are another example demonstrating the complex mechanisms by which climate change can cause damaging decreases in biodiversity and again provide an impetus to document the vast unknown biodiversity on earth in this case plant chemical diversity and insect diversity (see also Smilanich et al and Glassmire et al. 2016). We also made progress towards documenting this biodiversity and continued to discover new interactions, new phytochemicals, and new species of insects at all of our sites. Another paper that was more specifically about climate change used data on adult butterflies to demonstrate that climate change parameters have very different effects on insects, depending on the attributes of the species and site-specific attributes, such as unpredictable weather (Pardikes et al. 2015). Finally, we contributed a couple of papers to help clarify a debate about whether or not summer populations of monarch butterflies are in decline (Davis and Dyer 2015) it is still not clear what those populations are doing, but we do know that what is missing are data on the effects of changing plant chemistry, parasitoid diversity, and other natural enemies on these special butterflies (Dyer and Forister 3

4 2016). We continue to model the importance of interaction diversity as buffers to climate change and disruptive effects of losing biodiversity and we plan to continue the research over the coming decades. Figure 1. The concentrations and numbers of different plant toxins ("phytochemical diversity") causes greater herbivore diversity, greater plant toxicity, and lower herbivory in tropical rainforest shrubs ("pepper plants"). Positive causal relationships are shown in blue with arrowheads and negative relationships are indicated in red with bullet-heads. Plots of the partial correlations for each relationship are shown with the dependent variable on the x-axis and the response variable on the y-axis. Subsequent work by our collaborative team will examine how phytochemical diversity (Lewinsohn and Gijzen 2009) is affected by global change parameters. Our Earthwatch teams in Costa Rica and Ecuador collected data and specimens for this project, the Earthwatch teams in Nevada helped with the chemistry. Objective 2: Determine how hurricanes and variation in climate affect levels of parasitism and caterpillar densities. Models of climate change have predicted greater frequency and duration of droughts and floods and a widespread increase in the frequency of extreme weather events (e.g., Rahmstorf and Coumou2011). Such increased unpredictability and variability in regional climates, particularly with regard to precipitation, will likely impact interaction diversity, yet the potential effects 4

5 are largely unknown. As we reported in multiple previous Earthwatch reports, our hurricane damaged sites have experienced outbreaks by 8 different caterpillar species that reached outbreak proportions in response to storm damage. This is part of a general pattern for which outbreaks are more likely at sites which experience more variance in precipitation. For our Arizona and Sierra Nevada sites, one clear consequence of climate change and increased drought is that fires are increasing in intensity and frequency in arid regions and decreasing in other regions that are more fire adapted (Moritz et al. 2012). At our Florida and Sierra Nevada site, Earthwatch volunteers helped us examine the relationship between fire and plant and insect diversity. We have made progress at Eglin Air Force Base, which is the last extensive area of protected longleaf pine forest. We used Earthwatch plot data and previous diversity data to examine spatial patterns of diversity across Eglin, between forest types, and across a gradient along the Florida panhandle. As with all ecological relationships, our results are complex, but they provide insight into how plant diversity can be maintained by healthy fire intervals (Figure 2). We still have plenty of work to do to predict how insects and interactions are associated with different levels of fire. Figure 2. Summary of a structural equation model that shows relationships between fire, our measures of forest attributes, and understory plant diversity at our Florida Earthwatch site. Overstory, midstory, understory, and surface densities were quantified using satellite data (LIDAR). Positive effects are shown with blue headed arrows and negative correlations in red lines. Line thickness corresponds to strength of the relationship. Black dashed line represents total indirect effects of fire on diversity. Although the relationships are complex, the effects of fire on plant diversity are very clear and are likely to be affected by increases in drought. Earthwatch teams in Florida, Arizona, and the Sierra Nevada will continue to examine how plant chemistry, arthropods, and interactions are affected by fire frequency. 5

6 In addition to the modeling approaches mentioned in Objective 1, we continue to document the details of phenological asynchrony between parasitoids and their hosts when weather events such as floods and droughts create temporal shifts in caterpillar populations such that they are unavailable to specialized wasp parasitoids during their host-searching phase (Stireman et al. 2005). To this end, we hope to accumulate at least 2 more years of data to provide informationrich time series to allow us to determine which subfamilies and genera of parasitoids are most affected by specific extremes in weather. Objective 3: Specialization, trophic interactions, and biodiversity The diet breadth or host range of insects has been a focal point for understanding current levels of biodiversity, which are increasingly threatened by global change. Numbers of species that are published for the most diverse forests on earth are only crude approximations and the methods for estimating biodiversity rely on poor natural history data and untested assumptions about herbivore specialization (Basset et al. 2012). For years our Earthwatch data have facilitated investigation of specialization and co-diversification across multiple trophic levels, a long-standing challenge in biodiversity research and evolutionary theory. In 2015 we made excellent progress towards documenting specialization across climatic and latitudinal gradients, and we published 6 papers about this topic with a highlight being our paper on this topic in the prestigious journal, PNAS in which we report global similarities in distribution of diet breadth and a strong pattern of higher frequency of specialized insects in tropical regions (Forister et al. 2015). Our other papers explore the idea that our results on trophic interactions between insects and plants are relevant to other systems (e.g., Dyer et al. 2015, Rodriguez-Castaneda et al. 2016) for example at our Arizona site, we found that the negative effects of raptors on Mexican jay foraging cascaded down to increase hummingbird nest densities (Greeney et al. 2015; Figure 3). This result, published in the journal Science Advances, is very similar to many earlier Earthwatch-supported publications in which we reported such cascades for predator-herbivore-plant chemistry interactions. 6

7 Figure 3. Still not in love with caterpillars and parasitoids? Our results are also relevant to vertebrate communities! This figure depicts the cone-shaped space surrounding active hawk nests at SWRS Arizona. Hummingbird nests had significantly higher survivorship within this cone because hawks deterred jays (similar to the indirect positive effects of parasitoids on plants and plant chemistry). Yellow = hawk nest, Green = successful hummingbird nest, Red = hummingbird nest preyed on by jays. Objective 4: Discovery and descriptions of new species and new life histories As we have reported in past years, another significant accomplishment for 2015 came in the form of new species and new chemistry discoveries. For example, we reported on how three new chemical compounds (discovered by our team) at our Ecuador site are responsible for structuring populations of caterpillars and their associated parasitoids (Glassmire et al. 2016), which is a significant discovery. We have reported in many other papers that plant chemistry changes with increases in temperature and CO 2 (Dyer et al. 2012), and this paper shows similar changes across an elevational gradient. Chemical compounds produced by plants are not only responsible for the plants unique flavors, aromas, and colors, but also possible deterrent or toxic properties and resistance to herbivores (e.g., Hansen et al. 2016). Also similar to previous years, in 2015, we attempted to improve our sister site to caterpillars.org this is a site created in collaboration with the Encyclopedia of Life (EOL). The URL for the site is: Both of these 7

8 websites are in serious need of updates, and we plan to completely revamp them in the next two years, with the help of volunteers. In addition to updates to the life history data and images newly available on the web, we published several new life history descriptions for our focal taxa, many of which are the first biological information known for particular genera. We continue to rear out new species from the project at a rapid rate and we continue to publish species descriptions for as many of these as possible (Figure 4). At the Arizona site volunteers contributed to the Western North America caterpillar guide, the sequel to Dave Wagner s Eastern Caterpillars book. Figure 4. A geometrid caterpillar from the Arizona Earthwatch site. The juniper caterpillar and parasitoid communities were completely unknown before we started this project. In 2016 we will be getting sequences for all of the caterpillars we have reared and we will have the definitive networks for these communities in California, Nevada, and Arizona. A majority of these data will be new network connections. This type of discovery of new life histories and interactions is common at all of our sites. 8

9 PROJECT IMPACTS 1. Increasing Scientific Knowledge a. Total citizen science research hours - provide an estimate for the number of hours per day that volunteers spent collecting data, being trained to collect data in the field, and performing data entry. Volunteers typically work 8 hour days. This can be parsed between data collecting (4hrs), training during the first few days (4hrs), lab work (3-4hrs), and transportation (1hr). b. Peer-reviewed publications (2015/2016) Glassmire, A. E., Jeffrey, C. S., Forister, M. L., Parchman, T. L., Nice, C. C., Jahner, J. P., Wilson, J. S., Walla, T. R., Richards, L. A., Smilanich, A. M., Leonard, M. D., Morrison, C. R., Simbaña, W., Salagaje, L. A., Dodson, C. D., Miller, J. S., Tepe, E. J., Villamarin-Cortez, S. and Dyer, L. A. (2016), Intraspecific phytochemical variation shapes community and population structure for specialist caterpillars. New Phytol. doi: /nph Rodríguez-Castañeda, G., Brehm, G., Fiedler, K. and Dyer, L.A., Ant predation on herbivores through a multitrophic lens: how effects of ants on plant herbivore defense and natural enemies vary along temperature gradients. Current Opinion in Insect Science, 14: Hansen, A.C., A.E. Glassmire, L.A. Dyer, and A.M. Smilanich Cascading effects of diet induced variation in the immune response of two specialist herbivores. Ecography. Smilanich, A.M., R.M. Fincher, and L.A. Dyer Does plant-apparency matter? Insights from 30 years of data reveal clear patterns of the effects of plant chemistry on herbivores. New Phytologist 210: Scherrer, S., C. Lepesqueur, M.C. Vieira, M. Almeida-Neto, L.A. Dyer, M.L. Forister, I. Rezende Diniz Seasonal variation in host plant specialization by folivorous lepidopterans: contrasting patterns at the species and community levels. Biotropica. Dyer, L.A. and M.L. Forister Wherefore and whither the modeler: understanding the population dynamics of monarchs will require integrative and quantitative techniques. Annals of the Entomological Society of America. Richards, L.A., L. A. Dyer, M.L. Forister, A.M. Smilanich, C.D. Dodson, M.D. Leonard and C. S. Jeffrey Phytochemical diversity drives tropical plant-insect community diversity. Proceedings of the National Academy of Sciences 112: Greeney, H.F., R. Meneses, C.E. Hamilton, E.R. Hough, E.K. Austudillo, E. Lichter-Marck, R.W. Mannan, N. Snyder, H. Snyder, C.M. Ripplinger, S.M. Wethington, and L.A. Dyer Trait-mediated trophic cascade creates enemy-free space for nesting hummingbirds. Science Advances - 1, e Davis, A.K. and L.A. Dyer Long-term trends in eastern North American monarch butterflies: a collection of studies focusing on spring, summer and fall dynamics. Annals of the Entomological Society of America 108: Pardikes, N.A., A.M. Shapiro, L.A. Dyer, and M.L. Forister Global weather and local butterflies: variable responses to a large-scale climate pattern along an elevational gradient. Ecology 96: Forister, M. L., Novotny, V., Panorska, A. K., Baje, L., Basset, Y., Butterill, P. T., Cizek, L., Coley, P. D., Dem, F., Diniz, I. R., Drozd, P., Fox, M., Glassmire, A., Hazen, R., Hrcek, J., Jahner, 9

10 J. P., Kama, O., Kozubowski, T. J., Kursar, T. A., Lewis, O. T., Lill, J., Marquis, R. J., Miller, S. E., Morais, H. C., Murakami, M., Nickel, H., Pardikes, N., Ricklefs, R. E., Singer, M. S., Smilanich, A. M., Stireman, J. O., Villamarín-Cortez, S., Vodka, S., Volf, M., Wagner, D. L., Walla, T., Weiblen, G. D., and L. A. Dyer Global distribution of diet breadth in insect herbivores. Proceedings of the National Academy of Sciences 112: Dyer, L.A., T.J. Massad, and M.L. Forister The question of scale in trophic ecology. Pages in: Hanley, T. and K. La Pierre (eds.). Trophic Ecology: Bottom-Up and Top-Down Interactions across Aquatic and Terrestrial Systems. Cambridge University Press, Cambridge, MA. c. Non-peer reviewed publications: Technical reports, white papers, articles, sponsored or personal blogs d. Non-peer reviewed publications: Books and book chapters Dyer, L.A. and M.L. Forister The lives of Lepidopterists. Springer. e. Presentations : "Redes locais e globais em ecologia e evolução.", L. Dyer, invited departmental seminar, University of Campinas. (December 2015). "Climate change and interaction diversity", L. Dyer, N. Pardikes, Annual Meeting Entomological Society of America, Invited symposium talk, ESA. (November 2015). "Global weather and local butterflies: Variable responses to a large-scale climate phenomenon.", N. Pardikes, M. L. Forister, L. Dyer, A. Shapiro, regular talk, Annual Meeting Entomological Society of America, ESA. (November 2015). "A tropical story of natural history, Omics, and multi-trophic interaction diversity.", L. Dyer, invited departmental seminar, University of Utah. (October 2015). "Local versus regional networks of phytochemically mediated plant-caterpillar-parasitoid interactions", L. Dyer, invited symposium talk, 19th European Congress of Lepidopterology, Societas Europaea Lepidopterologica. (September 2015). "Intraspecific phytochemical variation: The ecological and evolutionary dynamics of chemical interactions.", A. E. Glassmire, C. Jeffrey, A. Smilanich, L. Richards, L. Dyer, et al., regular talk, Annual meeting, Ecological Society of America, ESA. (August 2015). "Quantifying the effects of burning on the diversity of arthropods in a fire-adapted longleaf pine (Pinus palustris) forest.", J. Dell, L. Richards, L. Dyer, et al., regular talk, Annual meeting, Ecological Society of America, ESA. (August 2015). Phytochemical diversity drives tropical plant-insect community diversity.", Richards, L. C. Jeffrey, A. Smilanich, L. Dyer, et al., invited seminar, Annual meeting, Phytochemical Society of North America. (August 2015). "Biodiversidad de Lepidoptera, Hymenoptera, y Diptera en los tropicos.", L. Dyer, Serra Bonita Biological Reserve. (March 2015). "Diversification in a tropical moth radiation: investigating the roles of phytochemistry, host history, and geography.", J. Jahner, L. A. Richards, T. L. Parchman, L. Dyer, A. Smilanich, C. Jeffrey, A. Glassmire, M. L. Forister, poster, Gordon Conference, Speciation. (March 2015). 10

11 2. Mentoring a. Graduate students list graduate students doing thesis work on the project Student Name Graduate Degree Project Title Anticipated Year of Completion Andrea Glassmire PhD Causes and consequences of intraspecific phytochemical variation on the physiology, abundance, and diversity of specialist consumers Nick Pardikes PhD Determinants of interaction diversity in Juniper woodlands Jane Dell PhD Untangling patterns of diversity in a fire forest: Spatiotemporal influence and interactions Danielle Salcido MS Global change, citizen science, and effective outreach for all ages Tara Langus MS Roles of pathogens, beneficial microbiota, and plant chemistry in the immune response of caterpillars b. Community outreach provide details on how you have supported the development of environmental leaders in the community in which you work. Name of school, organization, or group Education level Participants local or non-local Details on contributions/ activities Nevada Bugs and Butterflies Daugherty Summer Science Exploration All levels local Worked with Earthwatch volunteers and uses Earthwatch data as part of general outreach. K-9 local Worked with Earthwatch volunteers in the field. UNR Museum All levels local Earthwatch volunteers assisted with National Pollinator Week activities. 11

12 3. Partnerships list your current active professional partnerships that contribute to your project Partner Support Type(s) 1 Years of Association (e.g present) Serra Bonita Reserve, Bahia, Brazil Fundo Genova, Chanchamayo, Peru Organization for Tropical Studies Smithsonian National Museum of Natural History (UNMSM) American Museum of Natural History Museo de Historia Natural, Lima, Peru Museo Ecuatoriano de Ciencias Naturales (MECN) University of Nevada Natural History Museum University of Brasilia FAPESP Sao Paulo Research Foundation Conselho Nacional de Desenvolvimento Científico e Tecnológico Yanayacu Biological Station Southern Research Station, USFS partners in research, access to specimens, accepts voucher specimens in museum, contributes to conservation goals partners in research, access to specimens, contributes to conservation goals partners in research, access to specimens, contributes to conservation goals partners in research, provides space and access to specimens, accepts voucher specimens in museum partners in research, provides space and access to specimens, accepts voucher specimens in museum partners in research, provides space and access to specimens, accepts voucher specimens in museum partners in research, provides space and access to specimens, accepts voucher specimens in museum partners in research, provides space and access to specimens, accepts voucher specimens in museum partners in research, access to specimens provides additional funding and contributes to outreach goals provides additional funding and contributes to outreach and training goals (Science without Borders) partners in research, contributes to conservation and outreach goals provide funding for the Florida site, collaborate, contribute to management plans for Eglin 2014-present 2013-present 1991-present 2004-present 2004-present 2012-present 2002-present 2015-present 2008-present 2014-present 2013-present 2001-present 2012-present 12

13 4. Contributions to management plans or policies - list the management plans/policies to which your project contributed this year Plan/Policy Name Type 2 Level of Impact 3 New or Existing? Primary goal of plan/policy 4 Stage of plan/policy 5 Description of Contribution 2. Type options: agenda, convention, development plan, management plan, policy, or other (define) 3. Level of impact options: local, regional, national, international 4. Primary goal options: cultural conservation, land conservation, species conservation, natural resource conservation, other 5. Stage of plan/policy options: proposed, in progress, adopted, other (define) 5. Conserving natural and sociocultural capital a. Conservation of taxa i. List any focal study species that you did not list in your most recent proposal Species Common name IUCN Red List category Local/regional conservation status Local/regional conservation status source ii. In the past year, has your project helped conserve or restore populations of species of conservation significance? If so, please describe below. Species IUCN Red List category Local/regional conservation status Local/regional conservation status source Description of contribution Resulting effect 6 b. Conservation of ecosystems in the past year, has your project helped conserve or restore habitats? If so, please describe below. Habitat type Habitat significance 7 Description of contribution Resulting effect 8 Cloud forest Extremely high levels of diversity and endemism. Providing an economic incentive for managing lands for biodiversity by supporting the only research station in the area and providing longterm employment to local citizens. Yanayacu Biological Station and surrounding lands are still protected from logging, grazing, and development. 13

14 c. Ecosystem services Indicate which ecosystem service categories you are directly studying in your Earthwatch research and provide further details in the box below. Food and water Flood and disease control Spiritual, recreational, and cultural benefits Nutrient cycling Details: We continued to utilize a methodology that combined data using completely standardized methods in Costa Rica, Ecuador, Brazil, Argentina, Peru, and the United States to create plant-caterpillar-parasitoid diversity databases that can be used by land managers to search for natural enemies of agricultural pests and to examine relationships between enemy diversity and outbreak potential. Parasitoids provide an important, yet mostly unquantified ecosystem service that needs to be documented globally before it is enhanced, restored, or maintained. d. Conservation of cultural heritage provide details on intangible or tangible cultural heritage components that your project has conserved or restored in the past year. Cultural heritage Description of contribution Resulting effect component 9 9. Cultural heritage component options: traditional agriculture, artifacts, building(s), hunting ground or kill site, traditional ecological knowledge and practices, monument(s), oral traditions and history, spiritual site, traditional subsistence living Acknowledgements We first thank all of the amazing volunteers for hard work, patience, sharing their skills, and producing high quality ecological data and natural history observations. The support of Earthwatch Institute and the quality staff there is also greatly appreciated. This study was co-funded by NSF grants DEB , DEB , DEB , DEB , DEB , DEB , DEB , IOS ,the Department of Defense, and the University of Nevada. Our projects would not have functioned without expert assistance from many field technicians and colleagues who helped in the field and laboratory, gave lectures to volunteers, and discussed ideas with us; these include: M. Forister, T. Parchman, P. Poopat, C. Morrison, A. Palmer, M. Robinson, G. Blaustein, W. Lumpkin, J. Miller, S. Rab-Green, J. Jahner, R. Lindgren, A. Dietrick, R. Hazen, H. Greeney, D. Wagner, M. Singer, L. Salazar Vega, W. Simbaña, J. Simbaña, T. Walla, K. Fredrickson, T. Barber, H. Garcia Lopez, W. Simbaña, M. Ryan, M. Fox, T. Massad, and C. Pearson. 14

15 Literature Cited Davis, A.K. and L.A. Dyer Long-term trends in eastern North American monarch butterflies: a collection of studies focusing on spring, summer and fall dynamics. Annals of the Entomological Society of America 108: Dyer, L.A., Richards, L.A., Short, S. A., Dodson, C.D Effects of CO 2 and temperature on tritrophic interactions. PLOS ONE 8(4): e Dyer, L.A. and M.L. Forister Wherefore and whither the modeler: understanding the population dynamics of monarchs will require integrative and quantitative techniques. Annals of the Entomological Society of America. Dyer, L.A., T.J. Massad, and M.L. Forister The question of scale in trophic ecology. Pages in: Hanley, T. and K. La Pierre (eds.). Trophic Ecology: Bottom-Up and Top-Down Interactions across Aquatic and Terrestrial Systems. Cambridge University Press, Cambridge, MA. Forister, M. L., Novotny, V., Panorska, A. K., Baje, L., Basset, Y., Butterill, P. T., Cizek, L., Coley, P. D., Dem, F., Diniz, I. R., Drozd, P., Fox, M., Glassmire, A., Hazen, R., Hrcek, J., Jahner, J. P., Kama, O., Kozubowski, T. J., Kursar, T. A., Lewis, O. T., Lill, J., Marquis, R. J., Miller, S. E., Morais, H. C., Murakami, M., Nickel, H., Pardikes, N., Ricklefs, R. E., Singer, M. S., Smilanich, A. M., Stireman, J. O., Villamarín-Cortez, S., Vodka, S., Volf, M., Wagner, D. L., Walla, T., Weiblen, G. D., and L. A. Dyer Global distribution of diet breadth in insect herbivores. Proceedings of the National Academy of Sciences 112: Glassmire, A. E., Jeffrey, C. S., Forister, M. L., Parchman, T. L., Nice, C. C., Jahner, J. P., Wilson, J. S., Walla, T. R., Richards, L. A., Smilanich, A. M., Leonard, M. D., Morrison, C. R., Simbaña, W., Salagaje, L. A., Dodson, C. D., Miller, J. S., Tepe, E. J., Villamarin-Cortez, S. and Dyer, L. A. (2016), Intraspecific phytochemical variation shapes community and population structure for specialist caterpillars. New Phytol. doi: /nph Greeney, H.F., R. Meneses, C.E. Hamilton, E.R. Hough, E.K. Austudillo, E. Lichter-Marck, R.W. Mannan, N. Snyder, H. Snyder, C.M. Ripplinger, S.M. Wethington, and L.A. Dyer Trait-mediated trophic cascade creates enemy-free space for nesting hummingbirds. Science Advances - 1, e Hansen, A.C., A.E. Glassmire, L.A. Dyer, and A.M. Smilanich Cascading effects of diet induced variation in the immune response of two specialist herbivores. Ecography. Lewinsohn E, Gijzen M Phytochemical diversity: The sounds of silent metabolism. Plant Sci. 176: Moritz, M.A., Parisien, M.A., Batllori, E., Krawchuk, M.A., Van Dorn, J., Ganz, D.J. and Hayhoe, K., Climate change and disruptions to global fire activity. Ecosphere, 3:1-22. Pardikes, N.A., A.M. Shapiro, L.A. Dyer, and M.L. Forister Global weather and local butterflies: variable responses to a large-scale climate pattern along an elevational gradient. Ecology 96: Rahmstorf, S. and Coumou, D., Increase of extreme events in a warming world. Proceedings of the National Academy of Sciences, 108(44), pp Richards, L.A., L. A. Dyer, M.L. Forister, A.M. Smilanich, C.D. Dodson, M.D. Leonard and C. S. Jeffrey Phytochemical diversity drives tropical plant-insect community diversity. Proceedings of the National Academy of Sciences 112: Rodríguez-Castañeda, G., Brehm, G., Fiedler, K. and Dyer, L.A., Ant predation on herbivores through a multitrophic lens: how effects of ants on plant herbivore defense and natural enemies vary along temperature gradients. Current Opinion in Insect Science, 14: Scherrer, S., C. Lepesqueur, M.C. Vieira, M. Almeida-Neto, L.A. Dyer, M.L. Forister, I. Rezende Diniz Seasonal variation in host plant specialization by folivorous lepidopterans: 15

16 contrasting patterns at the species and community levels. Biotropica. Smilanich, A.M., R.M. Fincher, and L.A. Dyer Does plant-apparency matter? Insights from 30 years of data reveal clear patterns of the effects of plant chemistry on herbivores. New Phytologist 210: Stireman III, J.O., L.A. Dyer, D.H. Janzen, M.S. Singer, J.T. Lill, R.J. Marquis, R.E. Ricklefs, G.L. Gentry, W. Hallwachs, P.D. Coley, J.A. Barone, H.F. Greeney, H. Connahs, P. Barbosa, H.C. Morais, and I.R. Diniz Climatic unpredictability and caterpillar parasitism: implications of global warming. Proceedings of the National Academy of Sciences 102: