United States Department of Agriculture Proceedings of the American Elm Restoration Workshop 2016 Forest Service Northern Research Station General Technical Report NRS-P-174 September 2017 Abstract Proceedings from the 2016 American Elm Restoration Workshop in Lewis Center, OH. The published proceedings include 16 papers pertaining to elm pathogens, American elm ecology, and American elm reintroduction. This document is being published in electronic format only (Web). Any corrections or additions will be posted to the Website (https://doi.org/10.2737/NRS-GTR-P-174.) Cover Photo Baldwin Hill elm, late summer, 2013. Baldwin Hill crests between North and South Egremont in the southern Berkshires of western Massachusetts. The elm is growing on conservation farmland and was the first heritage American elm protected by the Elm Watch Adopt an Elm program. ©Tom Zetterstrom 2013, used with permission. The findings and conclusions of each article in this publication are those of the individual author(s) and do not necessarily represent the views of the U.S. Department of Agriculture or the Forest Service. All articles were received in digital format and were edited for uniform type and style. Each author is responsible for the accuracy and content of his or her paper. The use of trade, firm, or corporation names in this publication is for the information and convenience of the reader. Such use does not constitute an official endorsement or approval by the U.S. Department of Agriculture or the Forest Service of any product or service to the exclusion of others that may be suitable. This publication/database reports research involving pesticides. It does not contain recommendations for their use, nor does it imply that the uses discussed here have been registered. All uses of pesticides must be registered by appropriate State and/or Federal agencies before they can be recommended. CAUTION: PESTICIDES CAUTION: Pesticides can be injurious to humans, domestic animals, desirable plants, and fish or other wildlife—if they are not handled or applied properly. Use all pesticides selectively and carefully. Follow recommended practices for the disposal of surplus pesticides and pesticide containers. Manuscript received for publication April 2016 Published by U.S. FOREST SERVICE 11 CAMPUS BLVD SUITE 200 NEWTOWN SQUARE PA 19073 September 2017 Proceedings of the American Elm Restoration Workshop 2016 Lewis Center, OH October 25-27, 2016 Compiled and edited by: Cornelia C. Pinchot, U.S. Forest Service Kathleen S. Knight, U.S. Forest Service Linda M. Haugen, U.S. Forest Service Charles E. Flower, U.S. Forest Service James M. Slavicek, U.S. Forest Service Sponsored by: U.S. Forest Service, Northern Research Station General Technical Report NRS-P-174 Foreword On three beautiful fall days in central Ohio, an interdisciplinary group of scientists from across the United States and Canada came together at the American Elm Restoration Workshop to discuss past, present, and future research on American elm restoration (see page 144 for list of workshop participants). From October 25 to 27, 2016, the researchers met at Highbanks Metro Park in Lewis Center, OH, to capture historical insights into Dutch elm disease (DED) and the American elm, review the current status and evaluate the goals of future research, and foster collaboration among elm scientists. Of particular interest was garnering unpublished knowledge from scientists who were part of the “second front” of American elm research, circa 1960–1990, which benefited from substantial Federal funding. Capturing insights and knowledge from soon-to-be- and already-retired scientists was thus a primary goal of this workshop. The USDA Forest Service Northern Research Station and State & Private Forestry jointly hosted this workshop, with generous support from the Manton Foundation. The workshop covered a variety of topics relating to American elm restoration, including the DED pathogen, DED tolerance, genetics and ecology of the elm hosts, other threats to American elm, other tools to combat DED, and practical aspects of restoration. Together, the workshop presentations, partially represented by the papers in these proceedings, described challenges to American elm restoration, recent advances in the field, and research needs. As an introduction to the workshop, James M. Slavicek began with a description of an American elm restoration project involving the USDA Forest Service Northern Research Station and The Nature Conservancy. Some aspects of this project include screening survivor and progeny elms, represented in the paper by Charles E. Flower and colleagues (p. 24). Michael Marcotrigiano laid the groundwork for discussions with an overview of the research that has been done on DED and American elm, highlighting many of the answered and unanswered questions (p. 2). Louis Bernier helped us to focus briefly on the DED pathogen and warned of the potential of Ophiostoma spp. to overcome DED tolerance in DED-tolerant American elms, as well as the potential for the pathogen to jump to new insect vectors (p. 6). We proceeded to discuss the response of the elm host to DED through a series of talks on tolerance. Slavicek shared data that demonstrated heritability of DED tolerance. Crosses among DED-tolerant trees produced progeny with a range of tolerance phenotypes, including a large number of trees with excellent tolerance. Bernier provided information on several disease-response genes that are present in DED sensitive American elm but perhaps activated too late during pathogenesis to prevent disease. There are many traits that could be exploited in developing resistance, including anatomical features (size and spatial organization of vessels), timing of leaf flushing, preformed resistance molecules (phytoanticipins), induced resistance molecules (phytoalexins) and proteins, and attractiveness to elm bark beetles. Sherif M. Sherif shared data (published elsewhere) on activation of genes, demonstrating that tolerant American elms express certain genes more quickly and with greater magnitude than susceptible elms. Jasmonic acid is involved, and the response differs among tolerant elms, indicating multiple defense mechanisms. Alan T. Whittemore provided insight into the variable ploidy levels observed in American elm, alluding that the recently discovered diploid American elm that is more common in the southeastern United States may actually be a cryptic species and could provide a source of DED tolerance. Garrett L. Beier shared some preliminary results from his PhD research on compartmentalization response of American elms to inoculation with Ophiostoma novo-ulmi, demonstrating that some putatively tolerant clones show differential formation of zone 3 and zone 4 defense barriers. We followed these presentations with a panel discussion on methods used for challenge inoculations. Consistency and effectiveness in screening is necessary for accuracy and comparability of results. A summary of key observations from this panel are captured in these proceedings (Haugen et al, p. 37). Additional insights on how elm response to inoculation is affected by many factors, including time of year, age of tree, soil moisture, source and amount of inoculum, and method of delivery are provided in the papers by Beier et al. and Flower et al. in these proceedings (pgs. 30 and 24, respectively). We completed our topic on DED tolerance with a presentation by Tom Zetterstrom and colleagues about shortcomings in commercially-available elms, either in DED-tolerance or form (p. 119). Chad P. Giblin et al.’s paper provides further evidence of the failure of widely-planted cultivars in urban settings due to weak branch or leader attachments, unrelated to DED tolerance (p. 122). We also held an open discussion on the availability and characteristics of specific clones, as captured in the paper by Linda M. Haugen and Susan E. Bentz (p. 109). On the second day of the workshop, we resumed our consideration of the elm host with a series of talks more related to the importance of elms in the broader landscape. Jennifer L. Koch presented a summary of FIA data, which shows that in areas with longest presence of DED, some larger elms are persisting on the landscape. This persistence of large elms is not occurring in states where elm yellows is common, indicating that elm yellows may be contributing to the mortality of larger elms. Johanne Brunet found that the arrival of DED to North American has not reduced genetic diversity among slippery elm (Ulmus rubra), likely due to the high gene flow facilitated by wind-dispersed pollen and low levels of inbreeding, and suggested that DED is similarly unlikely to greatly diminish the genetic diversity of American elm (p. 99). Whittemore shared an analysis of the phylogeny of all known elm species, showing that our native North American elms (with the exception of Ulmus rubra) are in the subgenus Oreoptelea, whereas most of the “Old World” elms are in the subgenus Ulmus. It appears that several other species of elm could provide useful germplasm for breeding for resistance to DED and other valuable traits, particularly Chinese U. elongata and Himalayan U. villosa. As we shifted our consideration to other elm problems, Gary W. Moorman provided an overview of the elm yellows epidemic which occurred at Pennsylvania State University, and the lessons learned. There are three known strains of elm yellows: the European, common, and Illinois strains; the Pennsylvania outbreak was caused by the common strain. They observed that DED outbreaks often follow elm yellows, as the declining elms lead to increase in the elm bark beetles, which leads to an increase in DED. Cristina Rosa worked on identifying the insect vectors, and characterizing the elm yellows population, and her novel insights are captured in this proceedings (p. 49). In addition, Flower et al. recounted the complications of “false positives” that may be obtained using current molecular tools, because of presence of similar size and sequence DNA from other common bacteria (p. 68). Despite the many challenges to successfully growing American elm, there are useful tools to manage DED in high value trees.