Ts Denver Museum of Nature & Science Reports

DENVER MUSEUM OF NATURE & SCIENCE REPORTS DENVER MUSEUM OF NATURE & SCIENCE REPORTS DENVER MUSEUMDENVER OF NATURE & SCIENCE NUMBER 5, NOVEMBER 11, 2016

WWW.DMNS.ORG/SCIENCE/MUSEUM-PUBLICATIONS Denver Museum of Nature & Science Reports 2001 Colorado Boulevard (Print) ISSN 2374-7730 Denver, CO 80205, U.S.A. Denver Museum of Nature & Science Reports (Online) ISSN 2374-7749 REPORTS • NUMBER 5 NOVEMBER 11, 2016

Cover photo: Poladryas arachne (Nymphalidae) male, ventral; Kenosha Pass, Park County, Colorado, elevation 10,000 feet, July 27, 1985, collector: R. A. Watkins.

The Denver Museum of Nature & Science Reports (ISSN Frank Krell, PhD, Editor and Production 2374-7730 [print], ISSN 2374-7749 [online]) is an open- access, non peer-reviewed scientific journal publishing papers about DMNS research, collections, or other Program and Abstracts Museum related topics, generally authored or co-authored 27th Annual Meeting by Museum staff or associates. Peer review will only be arranged on request of the authors. of the High Country Lepidopterists

November 11 and 12, 2016 The journal is available online at www.dmns.org/Science/ Museum-Publications free of charge. Paper copies are University of Colorado Boulder exchanged via the DMNS Library exchange program ([email protected]) or are available for purchase from our print-on-demand publisher Lulu (www.lulu.com). M. Deane Bowers (Ed.) DMNS owns the copyright of the works published in the Reports, which are published under the Creative Commons Attribution Non-Commercial license. For commercial use of published material contact the Alfred M. Bailey Library & Archives at [email protected]. WWW.DMNS.ORG/SCIENCE/MUSEUM-PUBLICATIONS DENVER MUSEUM OF NATURE & SCIENCE REPORTS NUMBER 5, NOVEMBER 11, 2016

Program and Abstracts 27th Annual Meeting of the High Country Lepidopterists November 11 and 12, 2016, University of Colorado Boulder

Edited by CONTENTS M. Deane Bowers1 Program 2

Abstracts 3

27 Years High Country Lepidopterists’ Meetings 8

1University of Colorado Museum of Natural History and Department of Ecology and Evolutionary Biology UCB 334 University of Colorado Boulder, Colorado 80309, U.S.A. [email protected] Bowers 27th Annual Meeting of the High Country Lepidopterists

PROGRAM Friday, November 11, 6:00pm–9:00pm: Potluck dinner at the home of Deane Bowers, 10065 North 65th Street, Longmont, CO 80503 Saturday, November 12 9:00am–12:00pm: Open House in Entomology Section, CU Museum. Coffee and refreshments provided. The Entomology Section is on the 3rd floor of the MCOL Building. 12:00pm–1:30pm: Lunch (on your own; a group usually walks to a nearby café) 1:30pm–5:00pm: Contributed papers (Ramaley N240) 1:30pm: Deane Bowers: Welcome and introduction 1:35pm–1:50pm: Anna Sher, Seth Munson, Amerlia Bowman, Ryan Whitney, Francesca Aguirre-Wong, Eliot Jackson & Rob Robinson: Phenology of insect- pollinated high-altitude plants in Colorado 2:05pm–2:20pm: Ryan Beshai, Elizabeth Barnes & Shannon Murphy: The role of enemy- mediated competition in determining the fitness of a generalist herbivore 2:20pm–2:35pm: Elizabeth Barnes, Sarah Gosnell, Claudia Hallagan, Keelia Otten, Lainey Slayter & Shannon Murphy: Performance of western tent caterpillar (Malacosoma californicum) on two common host plants, including a new host plant record 2:35pm–2:50pm: Katherine Hernandez: Hommage à Trois: a tale of some very hungry Grammia spp. caterpillars 2:50pm–3:05pm: Paul Opler: The growing diversity of North American butterflies: what can we expect in the future? 3:05pm–3:30pm: Break 3:30pm–3:45pm: Jan Chu & David Ziskin: Trends emerge for butterfly populations in the Boulder County Open Space properties 3:45pm–4:00pm: Shannon Murphy, Mayra Vidal, Kylee Grenis, Whitley Lehto & Robin Tinghitella: Butterfly hunt: using a case study of butterfly predation to teach students about density dependence in Batesian and Müllerian mimicry 4:00pm–4:15pm: David Bettman: New and interesting microlepidoptera of Colorado 4:15pm–4:30pm: Chuck Harp: LepNet: a national project to digitize and image Lepido- ptera with contributions from the C.P. Gillette Museum, Colorado State University, Fort Collins 4:30pm–4:45pm: Timothy Smith: Effect of fire severity and age on plant-herbivore interactions 4:45pm–5:00pm: Mayra C. Vidal & Shannon M. Murphy: Influence of tri-trophic fitness on the diet breadth of a generalist herbivore 5:00pm: Business meeting 6:00pm: Dinner (at a local restaurant)

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The Role of Enemy-Mediated Competition ABSTRACTS—ORAL PRESENTATIONS in Determining the Fitness of a Generalist Herbivore

Performance of Western Tent Caterpil- Ryan Beshai, Elizabeth Barnes & Shannon Murphy lar (Malacosoma californicum) on Two Department of Biological Sciences, University of Common Host Plants, Including a New Denver, 2190 E. Iliff Avenue, Denver, Colorado 80208; Host Plant Record [email protected]

Elizabeth E. Barnes, Sarah Gosnell, Claudia Hallagan, The influence of competition on insect communities is Keelia E. Otten, Lainey Slayter & Shannon M. Murphy not well understood. We studied effect how the fitness Department of Biological Sciences, University of of a gregarious caterpillar species in Colorado, the Denver, 2190 E. Iliff Avenue, Denver, Colorado 80208; fall webworm (Hyphantria cunea), is affected by the [email protected] presence of another gregarious caterpillar species, the western tent caterpillar (Malacosoma americanum). Tent caterpillars are generalists across their full host Both tent caterpillars and fall webworms feed on range, but display local host plant preferences. We chokecherry as a host plant and create structures in present evidence for a new host plant record, wax which they reside until just prior to pupation. Prelimi- currant (Ribes cereum), for western tent caterpillars nary evidence suggests that tent caterpillar tents may (Malacosoma californicum) along the Colorado provide attractive physical and/or chemical cues for Front Range. We tested the suitability of wax currant as predators and parasitoids, thus increasing the enemy a host plant for western tent caterpillars as compared presence nearby the tents. We then tested whether tent to chokecherry (Prunus virginiana), an abundant caterpillar tents increase predation and parasitism and commonly used host plant. We measured the pressure on fall webworms colonies. density of tent caterpillar tents in areas where both host plants occur to assess host plant use. We reared tent caterpillar larvae on both host plants and mea- New and Interesting Microlepidoptera of sured fitness effects due to host plant quality (survival, Colorado pupal mass) and natural enemies (parasitism). We did not find a relationship between host plant abundance David Bettman and use by tent caterpillars and found no evidence Department of Zoology, Denver Museum of Nature & for a preference for either host plant. We found that Science, 2001 Colorado Boulevard, Denver, Colorado, western tent caterpillars do not differ in pupal mass 80205-5798; [email protected] when reared on chokecherry and on wax currant in a laboratory setting, but did vary in survival with greater The microlepidopteran fauna of Colorado is still far survival on wax currant. We found no difference in from fully explored. The primitive moth families Acan- parasitism rate for larvae collected from chokecherry thopteroctetidae (with two species on the Front Range, or wax currant. Our results suggest that wax currant is Acanthopteroctetes aurulenta and A. bimaculata) a suitable yet previously unrecorded host plant for tent and Eriocraniidae (with at least one new high-elevation caterpillar larvae. species) are here confirmed as new family records for the state. Many additional microlep taxa will be discussed and illustrated, documenting new state records, new life

history data, and more. of producing stacked images for high-resolution pho- Trends Emerge for Butterfly Populations in tographs. This presentation will give an overview of our the Boulder County Open Space Properties startup into this project and will give some early results.

Janet Chu & Daniel Ziskin 964 Ravenwood Road, Boulder, Colorado 80303; Hommage à Trois: a Tale of Some Very [email protected] Hungry Grammia spp. Caterpillars

For twelve years (2004–2016), the butterflies in eight Katherine Hernandez properties managed by Boulder County Parks and Open Department of Ecology and Evolutionary Biology, 334 Space (BCPOS) Colorado were inventoried. Although at UCB, University of Colorado, Boulder, Colorado 80309; the site with the highest elevation (Caribou Ranch) [email protected] some marker species, such as Queen Alexandra (Coleus alexandra), Rocky Mountain Dotted Blue (Euphilotes Larvae of the Arctiinae use an impressive arsenal of ancilla) and Small Wood Nymph (Cercyonis oetus), defenses to protect themselves against natural enemies. showed a slight increase in numbers over time, when Several species within this subfamily have been all eight properties and all species are combined studied for their defensive capabilities, specifically the there is a downward trend in butterfly abundance. In use of plant secondary metabolites. Life history and 2004, on average, a researcher could count about 42 sequestration patterns of secondary metabolites have butterflies per hour. Each year he or she would count been well documented for Grammia incorrupta; approximately one less per hour. By 2016, that same however, little information exists for other species of average researcher would count only 30. This paper Grammia. Furthermore, information on sequestra- will present the inventories by species and location and tion patterns during the immature stages of several sources of error for additional analysis. Grammia spp. is largely unknown. To address these gaps in knowledge, I conducted a comparative study of the chemical ecology and life history traits for the LepNet: a National Project to Digitize and virgin tiger moth (Grammia virgo), the little virgin Image Lepidoptera with Contributions tiger moth (Grammia virguncula) and the figured from the C. P. Gillette Museum, Colorado tiger moth (Grammia figurata) throughout larval State University, Fort Collins, Colorado development. I divided larvae into four treatment groups: one on white clover (Trifolium repens), one Chuck Harp on broadleaf plantain (Plantago major), one on 8834 W. Quarto Avenue, Littleton, Colorado 80128; narrowleaf plantain (Plantago lanceolata), and one [email protected] on a wheat-germ based artificial diet. I collected data on development time for each instar, larval weight at In the early summer of 2016, the Gillette Museum was each instar and overall survivorship. A subset of larvae awarded a four-year National Science Foundation Grant was also harvested at each instar; these samples were to participate in the LepNet Project. With 27 institutions prepared for chemical analysis to determine presence participating in the project, CSU’s Gillette Museum will or absence of aucubin and catalpol at different instars. database 156,000 butterflies and moths from its holdings Aucubin and catalpol are found in one or both of the and will photograph nearly 11,000 of these specimens Plantago species used in this study. Here I will present using a state-of-the-art digital camera system, capable results of the feeding trials and the chemical analyses

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of the Grammia tiger moths and discuss implications facilitate students’ understanding of such an involved of these results on the understanding the chemical topic. We designed an interactive activity in which stu- ecology and fitness of these species. dents learn about density dependence by experiencing how variation in density of palatable and noxious butterflies affects a predator’s ability to learn warning signals Colorado Butterfly Monitoring Network and thus the prey’s ability to avoid predation. In class, (Update) students act as a predator and “hunt” butterflies that are either palatable or noxious. As part of our butterfly case Morgan Kerr study, the students learn to distinguish between negative 9840 Westcliff Pkwy #1122, Westminster, Colorado and positive density dependence, graph and interpret 80021; [email protected] data, and how predation can lead to convergent evolu- tion in warning signals. Colorado has joined 12 other states and regions across the country in utilizing citizen scientists to monitor butterfly populations. The goal of these monitoring efforts The Growing Diversity of North American is to track butterfly distributions, timing, yearly changes, Butterflies: What Can we Expect in the unusual events, and dynamics at the local, regional, and Future? continental scale. The summer of 2013 was the pilot year for Colorado Butterfly Monitoring Network and included 8 Paul A. Opler volunteers who monitored 9 sites. After the first year, vol- P. O. Box 2227, Loveland, Colorado 80539 unteer efforts increased to include over 50 individuals who [email protected] monitored 46 sites. In 2016, the program now supports 166 active volunteers and 96 sites. The combined results from Historically, the number of butterfly species recognized the first three seasons include the identification of 111 in the North American fauna has grown at a more or species out of 23,557 observed butterflies. less constant rate of 2–4 species per year. Most recently the Pelham listed published in 2009 listed 800 species, but now the list has grown to about 814 species. This is Butterfly Hunt: Using a Case Study of But- due to the detection of species from Latin America being terfly Predation to Teach Students about detected along our southern borders and the discovery of Density Dependence in Batesian and Mül- new species complexes of what were previously thought lerian Mimicry to be single species through research into mitochondrial DNA, or more recently, the entire DNA complement Shannon Murphy, Mayra Vidal, Kylee Grenis, Whitley through genomics. Recent examples include splitting Lehto & Robin Tinghitella the Giant Swallowtail, Heraclides cresphontes, into two Department of Biological Sciences, University of species, and the recognition that the Carolina Satyr, Her- Denver, 2190 E. Iliff Avenue, Denver, Colorado 80208; meuptychia sosybius, also includes two cryptic sibling [email protected] species. We can expect this increased diversity to continue into the indefinite future. Some examples within Colorado Students usually do not completely understand the include cryptic species within the Sara Orangetip, Pine concept of density dependency in ecology due to the White, Shasta Blue, and Ridings Satyr to name a few. As complexity of the subject. Teaching cases of density a result of this process I expect that the North American dependence featuring classic evolutionary and ecological butterfly fauna will reach 850 by mid-century and more processes, such as Batesian and Müllerian mimicry, may than 900 before the beginning of the next millennium.

Phenology of Insect-Pollinated High- and Federally Listed rare/endangered Colorado species, Altitude Plants in Colorado but with differences between habitats. Surprisingly, the highest altitude species had the smallest shifts Anna A. Sher over time, with the greatest acceleration observed for Department of Biological Sciences, University of species in montane sagebrush interbasins. High winter Denver, Denver, Colorado 80208; [email protected] temperatures were associated with the acceleration of phenology in lower elevation habitats, whereas high Seth Munson spring temperatures explained accelerated phenology U.S. Geological Survey, Southwest Biological Science in the high elevation alpine habitat. Precipitation had Center, 2255 N. Gemini Dr., Flagstaff, Arizona 86001 mixed effects depending on the season. These findings suggest that flowering plants are highly sensitive to the Amelia Bowman, Ryan Whitney, Francesca AguirreWong, climate changes observed in high altitudes over the Department of Biological Sciences, University of past century, with changes more than twice as large as Denver, Denver, Colorado 80208 those observed on the East Coast. It is critical that these phonological shifts be compared against those of their Eliot Jackson pollinators if we are to understand the future of these St. Mary’s College of California, Moraga, California species in the context of climate change. 94575

Rob Robinson Effect of Fire Severity and Age on Plant- Research and Conservation, Denver Botanic Gardens Herbivore Interactions 1007 York Street, Denver, Coloradio 80206 Timothy Smith Increasing temperatures across the Southern Rocky Department of Biological Sciences, University of Mountains have the potential to alter the phenology Denver, 2190 E. Iliff Avenue, Denver, Colorado of flowering plants and thus shift pollinator relation- 80208; [email protected] ships. Rare plants may be especially at risk given their isolated distributions, yet previous research had not Climate change models have predicted an increase investigated either the role of rarity or elevation in in the area burned by fires as well as the severity of the magnitude of change and therefore degree of risk. these fires across much of the US. Considering these Thus, we asked the following questions: 1) to what predictions, it is important to investigate the effects degree are species in the Southern Rocky Mountains that fire can have on forest ecosystems. Our study showing earlier flowering times and are these patterns tested how fire severity and the amount of time since seen in rare species? 2) To what degree are these shifts the fire affect herbivore damage on the plant wax related to measures of climate change? 3) Are these currant (Ribes cereum). We measured overall herbi- patterns consistent across habitat types and elevations? vore damage, but were particularly interested in plant To address these questions, we amassed label data from damage by the hoary comma butterfly Polygonia( over 600 flowering herbarium specimens of Colorado gracilis zephyrus), which leaves a unique stripping plants collected from 1870 to 2009 from herbaria pattern when it feeds. We surveyed plants from three across the country. We found these species to be flower- separate fires within Pike National Forest in Colo- ing approximately 20 days earlier now versus a century rado. For each fire, we selected 9 patches including ago. A significant negative correlation was found 3 high severity patches, 3 low severity patches, and between collection date and year for both common 3 unburned patches. In each patch, we surveyed 5

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plants for herbivore damage and noted if the damage parasitism rate and plant quality vary among popula- was due to Polygonia. In addition, we surveyed the tions. We found that the variation in top-down and density of wax currant at each patch by counting bottom-up forces led to different diet breadths; popula- the number of plants within a 1250m2 area. Using tions tend to be more specialized when they are already this data, we concluded that the amount of overall using the host plant species associated with greater tri- herbivore damage was significantly greater in low and trophic fitness, whereas populations using host plants unburned patches than in high severity patches. For associated with lower tri-trophic fitness had broader damage by Polygonia, however, we found significantly diets. Our results suggest that tri-trophic fitness is an more damage in the unburned patches than in the important selective force shaping diet breadth. low and high severity patches, which suggests that Polygonia butterflies fare better in unburned forests compared to forests recovering from forest fire.

Influence of Tri-Trophic Fitness on the Diet Breadth of a Generalist Herbivore

Mayra C. Vidal & Shannon M. Murphy Department of Biological Sciences, University of Denver, 2190 E. Iliff Avenue, Denver, Colorado 80208; [email protected]

The great majority of insect herbivores are specialists, however what drives specialization is not completely understood. Most research to understand specialization and speciation in herbivorous insects was done using specialists as study organisms. However, by studying why generalist species may be locally specialized in some populations but not others, we may better understand the drivers of specialization and of divergent selection among populations. We developed a conceptual framework including all possible scenarios of tri-trophic fitness associated with frequency of host plant use to predict the diet breadth of each herbivore population in a given time. Using a generalist herbivore (fall webworm, Hyphantria cunea, Erebidae; hereon FWW) with varied diet breadths across its wide geographic range, we are testing how top-down and bottom-up forces vary among populations of FWW and how those selective forces influence FWW diet breadth in the different populations. We used plant quality and parasitism rate as our bottom-up and top-down forces because FWW are greatly affected by them and both

1990: 1st meeting (High Plains Lepidopterists), September 14–15, Holiday Inn University Park, hosted by Colorado State University, Entomology Department, Fort Collins 1991: 2nd meeting (High Plains Lepidopterists), October 4–5, University of Colorado Museum, Boulder 1992: 3rd meeting, September 11–12, Denver Museum of Natural History, Denver 1993: 4th meeting, September 18, University of Wyoming, Department of Entomology Insect Collection, Laramie 1994: 5th meeting, October 28–29, Holiday Inn University Park, hosted by C.P. Gillette Museum of Arthropod Diversity Colorado State University, Fort Collins 1995: 6th meeting, October 20–22, University of Colorado Museum, Boulder 1996: 7th meeting, October 25–26, Butterfly Pavilion, Westminster 1997: 8th meeting, September 19–20, Holiday Inn University Park, Fort Collins 1998: 9th meeting, October 23–24, University of Colorado Museum, Boulder 1999: 10th meeting, October 22–23, C.P. Gillette Museum, Colorado State University, Fort Collins 2000: 11th meeting, November 3–4, C.P. Gillette Museum, Colorado State University, Fort Collins 2001: 12th meeting, September 7–8, C.P. Gillette Museum, Colorado State University, Fort Collins 2002: 13th meeting, October 11–12, University of Colorado Museum, Boulder 2003: 14th meeting, November 7–8, C.P. Gillette Museum, Colorado State University, Fort Collins 2004: 15th meeting, November 5–6, University of Colorado Museum, Boulder 2005: 16th meeting, October 21–22, C.P. Gillette Museum, Colorado State University, Fort Collins 2006: 17th meeting, October 27–28, University of Colorado Museum, Boulder 2007: 18th meeting, November 2–3, C.P. Gillette Museum, Colorado State University, Fort Collins 2008: 19th meeting, October 24–25, University of Colorado Museum, Boulder 2009: 20th meeting, October 23–24, Denver Museum of Nature & Science, Denver 2010: 21st meeting, November 5–6, C.P. Gillette Museum, Colorado State University, Fort Collins 2011: 22nd meeting, October 14–15, University of Colorado Museum, Boulder 2012: 23rd meeting, October 20, Butterfly Pavilion, Westminster 2013: 24th meeting, November 1–2, C.P. Gillette Museum, Colorado State University, Fort Collins 2014: 25th meeting, November 7–8, Denver Museum of Nature & Science, Denver 2015: 26th meeting, October 23–24, University of Denver, Denver 2016: 27th meeting, November 11–12, University of Colorado, Boulder

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Cover photo: Poladryas arachne (Nymphalidae) male, ventral; Kenosha Pass, Park County, Colorado, elevation 10,000 feet, July 27, 1985, collector: R. A. Watkins.