Department of Agriculture

Technology Transfer Non-native Pest

Biology and Control of Emerald Ash Borer

Edited by Roy G. Van Driesche and Richard C. Reardon

Forest Health Technology Enterprise Forest Service Team Morgantown, WV FHTET-2014-09 March 2015 The Forest Health Technology Enterprise Team (FHTET) was created in 1995 by the Deputy Chief for State and Private Forestry, USDA Forest Service, to develop and deliver technologies to protect and improve the health of American forests. This book was published by FHTET as part of the technology transfer series. http://www.fs.fed.us/foresthealth/technology/

On the cover: Cover design by Sheryl Romero and Denise Binion, Forest Health Technology Enterprise Team. Background image: Understory green ash seedlings ( pennsylvanica, ) released after large ash trees were killed by emerald ash borer in Okemos, Michigan in 2014, photo by Leah S. Bauer; (bottom row, left to right) Fully mature Tetrastrichus planipennisi larvae break free of emerald ash borer larval skin and pupate in the larval gallery under the tree bark. (Photo credit: Clifford Sadof); EAB adult and typical leaf feeding damage. (Photo credit: Deborah Miller, USDA Forest Service, Bugwood.org); Emerging Tetrastrichus plannipennisi adults. (Photo credit Leah S. Bauer).

We thank the authors of the individual chapters for their expertise in reviewing and summarizing the literature and providing current information on the biology and control of emerald ash borer. Thanks to Denise Binion for layout and design of this publication. We would also like to thank the U. S. Department of Agriculture, Forest Service, Agricultural Research Service and and Health Inspection Service for technical and financial support and Forest Health Technology Enterprise Team for providing funding for the preparation and printing of this publication.

For additional copies of this publication, contact: Richard Reardon Roy Van Dreische USDA Forest Service Department of Environmental Conservation 180 Canfield St. Holdsworth Hall Morgantown, WV 26505 University of Massachusetts, Amherst, MA 01003 (304) 285-1566 (413) 545-1061 [email protected] [email protected]

The entire publication is available online at:

http://www.fs.fed.us/foresthealth/technology/pdfs/FHTET-2014-09_Biology_Control_EAB.pdf

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Edited by

Roy G. Van Driesche

Richard C. Reardon

FHTET-2014-09 March 2015 Chapter 2: eCologiCal impaCts oF emerald ash Borer

David L. Wagner1 and Katherine J. Todd2

1Corresponding author, Center for Conservation and Biodiversity, Department of Ecology and Evolutionary Biology, University Box 43, University of Connecticut, Connecticut 06269, [email protected] (860)486-2139 2Department of Entomology, The Ohio State University, Columbus, Ohio 43210, [email protected]

introdUCtion and is expanding its range on all edges of its current distribution. Because timber and firewood from The genus Fraxinus includes 16 North American infested areas can be transported long distances, the . Flowers et al. (2013) estimate that there are has the potential to establish virtually anywhere more than 8.7 billion ash trees and saplings in the where ash species grow. It is considered to be among continental United States, and these are all potentially the most destructive forest pests to have susceptible to emerald ash borer (EAB) ( been introduced into North America (Herms and planipennis Fairmaire) infestation. Since its invasion, McCullough, 2014; McCullough and Usborne, 2014). this beetle has established in more than 20 states, More than 200,000 million ash trees have been killed from Minnesota to Maine, south to Georgia and – especially in the Great Lakes region where the Missouri, as well as in southern Ontario and Quebec, insect first established in the early 1990s (Cappaert et al., 2005a; Poland and McCullough, 2006, Siegert et

1 2

3 4

Figures 1-4. Affected communities in Ohio; dead trees are green ash F.( pennsylvanica) killed by the emerald ash borer: (1) Saint Mary’s River west of Decatur, June 2014. (Photo courtesy Jim McCormac); (2) Willow Point Wildlife Area in Vickery, August 2011. (Photo courtesy Jim McCormac); (3, 4) Ottawa National Wildlife Refuge in Oak Harbor, August 2014. (Photos courtesy Judy Semroc)

BIOLOGY AND CONTROL OF EMERALD ASH BORER 15 CHAPTER 2: ECOLOGICAL IMPACTS OF EMERALD ASH BORER

al., 2007) (Figs. 1-4). (Entomology Today, 2014). L. (white ash) and especially Climate models for the beetle and projections Fraxinus pennsylvanica Marshall (green ash) – strong, for its spread are still in development and burdened stately, rapidly growing trees – are widely planted in with considerable uncertainty (Sobek-Swant et al., yards, parks, and along city streets, in part because 2012; Liang and Fei, 2014), thus it is impossible both species are known to be hardy and relatively to know how far north, south, and west the beetle insect- and disease-free (Burns and Honkala, 1990). might spread in North America and which Fraxinus Ash is also preferred for firewood because its sap populations are most at risk. Consequently, for the is flammable and, as such, the potential for EAB to purposes of this assessment, we consider the entirety be spread throughout North America by human of the North American ash flora to be vulnerable. transport of infested wood is greatly heightened. We first examine the forest community types where A lower bound economic estimate for treating, ash is a dominant or co-dominant and then provide removing, and replacing EAB-infested ash trees in a brief assessment for each of the sixteen Fraxinus urban landscapes for 2010 to 2020 is $12.5 billion that grow north of Mexico. Each treatment includes (Kovacs et al., 2011). White ash and, to a lesser extent, a synopsis of a species’ habitat, range, conservation green ash are commercially important trees whose status, known susceptibility to EAB, and a summary strong but flexible wood is used for flooring, paneling, of its specialist herbivores. We then include a brief furniture, tool handles, and baseball bats (Elias, 1987; discussion of the importance of ash to vertebrate Burns and Honkala, 1990). White ash has long been wildlife before addressing the core of our contribution the preferred wood for the Louisville Slugger – a bat – a comprehensive evaluation of the invertebrate popular with professional baseball players. herbivore fauna of North American Fraxinus Ash typically grows as a component in hardwood likely to be threatened by the spread of EAB. Our forests (Figs. 5-11); furthermore, Fraxinus species evaluation is constructed from reviews of literature, are classified as either the dominant or co-dominant correspondence with taxonomic authorities, and species in 150 forest and shrubland communities DLW’s 30-year rearing program. We ignore generalist (NatureServe Explorer, 2014, see below). In some herbivores known to feed on ash, although a few wetlands, species such as Marshall oligophagous species are discussed. (black ash) (Figs. 8, 10) and Fraxinus profunda Our assessment is novel in that we considered (Bush) Bush (pumpkin ash) (Figs. 7, 9) form almost feeding records in older and derivative literature pure monocultures that are highly susceptible to as unconfirmed, given the large number of EAB (Tardif and Bergeron, 1992; Rebek et al., 2008; misattributed records plaguing recent compendia and Klooster et al., 2014; NatureServe Explorer, 2014; Jim risk-assessment literature. Instead, we adopted an McCormac and Jim Bissell pers. comms). Laboratory authority-driven approach whereby we contacted one trials suggest that all 16 species of North American or more active systematists or experts for taxa known Fraxinus may be suitable hosts for larval development to have specialist herbivores on woody plant taxa. We of EAB and thus are susceptible to attack (Anulewicz received information from more than 80 taxonomic et al., 2008; Leah Bauer pers. comm, Deb McCullough authorities with first-hand knowledge of appropriate pers. comm). Even small-stemmed western species literature, species-level , life history data, have the potential to support the beetle given that and ecological associations. Compared to previous saplings of white and green ash as narrow as 1 cm in assessments, our approach yielded a dramatically diameter are exploited by the beetle and its immature different list of potentially imperiled herbivores, stages in the . Once infested, tree essentially half of which are newly reported here. death typically follows within 2 to 6 years (Knight et al., Our work includes a revised set of risk rankings for 2013). In addition, the related (olive family) fringetree 98 species that we believe to be threatened by the ( L.) is also susceptible to attack, emerald ash borer. but the beetle’s impact on that host is as yet unstudied

BIOLOGY AND CONTROL OF EMERALD ASH BORER 16 CHAPTER 2: ECOLOGICAL IMPACTS OF EMERALD ASH BORER

5 6 7

8 9

10 11

Figures 5-11. Ash trees and ash-dominated communities: (5) Google Earth view of black ash swamp west of Lincoln, Wisconsin; Fraxinus nigra is dominant over thousands of hectares of this boreal wetland; (6) Majestic blue ash (F. quandrangulata) tree near Lexington, Kentucky. (Photo courtesy Daniel Boone); (7) Vernal pool with canopy of pumpkin ash (F. profunda), Momence Wetlands Land and Water Reserve, ; note buttress roots or knees which help the trees uptake oxygen when the pool is inundated. (Photo courtesy Judy Semroc); (8) Black ash dominated swamp near Black Lake, Michigan. (Photo courtesy Anton Reznicek); (9) Vernal pool near Astabula, Ohio with pumpkin ash as the dominant canopy tree; the dry vernal pool shown here is a breeding area for mole salamanders. (Photo courtesy Judy Semroc)—see Effects to Vertebrates; (10) Northern hardwood swamp dominated by black ash near Wallon Lake, Michigan. (Photo courtesy Anton Reznicek); (11) Upland hickory-white ash glade (F. americana) in Litchfield County, Connecticut. (Photo courtesy Ken Metzler)

BIOLOGY AND CONTROL OF EMERALD ASH BORER 17 CHAPTER 2: ECOLOGICAL IMPACTS OF EMERALD ASH BORER

methods and rationale USNVC classification. Ash specialists were defined as those species for To determine ecological impacts of EAB infestations which Fraxinus was deemed a principal larval or on communities, we contacted 20 people with adult host. Stated differently, specialists are those taxa first-hand experience (botanists, reserve managers, that would be expected to be severely compromised Department of Natural Resources (DNR) ecologists, were Fraxinus eliminated from the taxon’s range. State Nature Conservancy chapters, forest managers, We excluded more than 180 ash-feeding herbivores land managers, seasoned naturalists, wildlife that are known to feed on hosts outside the family biologists, and especially those with extensive field Oleaceae (Gandhi and Herms, 2010; Robinson et experience) in the forests and woodlands where EAB al., 2014). Host records were verified by contacting has been resident for four or more years (Appendix principal taxonomic experts (listed with affiliations 1). We focused our surveys and correspondence on in Appendix 2). Frequently, multiple experts, with three states and one province with conspicuous EAB differing regional data or taxonomic knowledge, were impacts: Michigan, northern Ohio, northeastern consulted for the same taxon. We were not able to , and southwestern Ontario. For each state make contact with an active North American thrips or province we initially contacted a lead biologist in (Thysanoptera) systematist and caution that much the DNR (or its functional equivalent) and a state or remains to be learned about North America’s provincial office for The Nature Conservancy and fauna. Documentation for all taxa and the listed hosts then made efforts to approach additional state or is given in the References column and, as appropriate, provincial authorities that had been endorsed for in the Comments column of Table 4. Conversations their knowledge of the impacts of EAB. Vertebrate are referenced as “pers. comm.” and letters and emails biologists that we contacted for information on the are referenced as “in litt.” ecological (especially dietary) importance of Fraxinus We use the term polyphagous to refer to cases are listed in Appendix 1. Conversations with Anton where herbivores feed on members of more than Reznicek (University of Michigan), renowned Great two families, oligophagous to refer to cases where Lakes Region botanist, Jim Bissell (Cleveland Museum the herbivores feed on more than one genus in of Natural Science), and Jim McCormac (Ohio the Oleaceae, and specialist to refer to cases where 1 Division of Wildlife) carried special force. Fraxinus is the sole or principal host in nature . We To identify and assess imperilment of plant restrict use of monophagous to those instances where communities containing ash, we used the U.S. only a single species is (known to be) consumed, National Vegetation Classification (USNVC) revised and ecological monophagy to instances where a in 2008 and subsequently developed by NatureServe species’ diet is restricted to a single member of Explorer and state Heritage Programs (NatureServe the Oleaceae because only one host species grows Explorer, 2014). The classification system for in a given geographic location; presumably the the North America’s plant community types – a herbivore would use other congeners (Fraxinus) or jurisdictional subset of the International Vegetation perhaps confamilials (Oleaceae) were these available. Classification – is a spatially extensive, range-wide, Throughout this paper, our diet-breadth assessment on-going collaboration of federal, international, of invertebrates applies to the most specialized life academic, and state partners, housed and managed stage; thus, if a eats just Fraxinus as a larva by NatureServe. Our treatment focuses on ecological but browses on a spectrum of as an adult, our communities recognized at the Group level in the discussion and risk assessment is based on the larval International Vegetation Classification system and stage. tallies the biological communities in which any one of North America’s 16 Fraxinus species is named as a 1This is more restrictive than most insect-plant literature, where oligophagous refers to herbivores that consume plants from dominant or co-dominant plant species. We capitalize just 2-3 families and polyphagous is used to refer to herbivores Group when the word is meant to convey rank in the consuming four or more families.

BIOLOGY AND CONTROL OF EMERALD ASH BORER 18 CHAPTER 2: ECOLOGICAL IMPACTS OF EMERALD ASH BORER

We follow Hinsinger et al. (2013) in recognizing herbivores also reported from non-native Oleaceae four Sections of Fraxinus in North America. Section hosts, e.g., Olea L. (olive), Ligustrum L. (), is capitalized when the word is meant to convey and L. (lilac) were given a High to Moderate nomenclatural rank (and phylogenetic membership), risk rating as these hosts are either considered not i.e., roughly equivalent to a subgenus. Authors for abundant enough in wildlands or too infrequently scientific names of imperiled are given used to serve as viable (sole) hosts for these taxa over in Table 4; author names for species not extended time periods were ash to be functionally treated in Table 4 are given in the text the first time eliminated. If a species was believed to have a strong that taxon is discussed. Arthropod family names are preference for Fraxinus over other Oleaceae, its given in the text and Table 4 except where family risk rating was increased; conversely, those with a membership has been made obvious by accompanying preference for Forestiera resulted in a reduced rating. text. Authors for Fraxinus species appear in the section One species, Prociphilus americanus (Aphididae), entitled North America’s Sixteen Fraxinus. was determined as High risk despite feeding on Abies Species-level taxa (n = 98) were assigned an Miller (fir) because both Fraxinus and fir are primary imperilment or risk category after we had verified host hosts in different stages of this aphid’s alternating life records for each. Four categories of risk were adopted: cycle. Very High, High, Moderate, and Low, with all examples In a few cases where we suspected an apparent of the latter excluded from this work. If only Fraxinus specialist may prove to be a polyphage, e.g., Banasa and native Chionanthus L. (fringetree) species were rolstonii (Pentatomidae), Diaspidiotus fraxini, and among the reliably reported hosts, individuals were Diaspis fraxini (both Diaspididae), we invoked the given a High risk rating. If additional Oleaceae hosts precautionary principle and assigned these species were recorded, we assigned risk ratings in the follow a High risk rating. Likewise, species in need of manner: herbivores known to include Forestiera Poir. taxonomic study were still included and generally (swamp privet or desert olive) species were regarded ranked as High; e.g., near chersis, Sympistis as Moderate risk species because native Forestiera fortis, Hyrdelia near inornata, and Zelleria near are abundant enough to serve as an alternative host hepariella. In these cases and others, our thinking is to Fraxinus; ash-feeding and fringetree-feeding conveyed in the Comments section of Table 4.

Figure 12. Google Earth image of Saint Mary’s River west of Decatur, Ohio. The dead trees in this image about the periphery in the riparian corridor east of Rockford are essentially all green ash. The red star indicates the approximate location of the stand shown in Figure 1.

BIOLOGY AND CONTROL OF EMERALD ASH BORER 19 CHAPTER 2: ECOLOGICAL IMPACTS OF EMERALD ASH BORER

ECOLOGICAL IMPACTS TO NORTH tree and the assemblage is regarded as either critically AMERICAN ASH imperiled (G1) or imperiled (G2) in NatureServe Explorer (2014). Most of these imperiled forest types Flowers et al. (2013) estimate that there are 8.7 billion occur south of Pennsylvania; three are restricted to ash trees and saplings in the lower 48 states – making Oregon and Washington. Fortunately, none occur in up roughly 2.5% of the aboveground forest carbon the most severely affected areas of the Midwestern mass in this region. Over most of North America, ash United States. trees tend to grow in mixed hardwood woodlands A few community types warrant special mention and forests and are infrequently an ecologically because of their spatial extent or because of their dominant tree at landscape levels. Even in regions vulnerability. In Wisconsin and elsewhere, swamps of the Midwestern United States, where a significant with black ash as the sole dominant canopy species portion of all Fraxinus have been killed, we were may encompass >1000 ha (Fig. 5). Palik et al. (2012) unable to easily detect affected communities using speculated that many such black ash swamps, should Google Earth satellite imagery; although one such ash disappear, may change permanently to shrublands example – a severely damaged area along the Saint as no other tree species exists in large enough Mary’s River, east of Decatur, Ohio – is shown in concentrations to form a new canopy. Similarly, along Fig. 12. Another important exception are black ash the Atlantic Coastal Plain, Carolina ash sometimes forests, which can be ecologically dominant at larger accounts for much of the above-ground biomass in spatial scales; in many northern woodlands, F. nigra bottomlands and wetlands. grows in virtual monocultures over hundreds of The most significant ecological impacts of EAB hectares (see below and Figs. 5, 8, 10). Furthermore, to woodlands and forests will be determined by what over smaller spatial scales, ash species have great plant associations establish post-invasion (Flowers et ecological importance and, by definition, are essential al., 2013; Knight et al., 2013; Burr and McCullough, elements in the woodlands and forests where they are 2014). Future projections for communities formerly dominant or co-dominant species. The U.S. National dominated by ash are still a matter of conjecture Vegetation Classification and NatureServe Explorer because no EAB infestations, with their concomitant (2014) identify 150 U.S. and Canadian forest and ecological consequences, have run full course. shrubland community types where a Fraxinus species Undoubtedly, there will be taxonomic variation in is named as a dominant or co-dominant element. which species replace ash due to differences in soil Just four Fraxinus species account for 82% of the 150 type, hydrology, light, seed banks, and the local pools community Groups where ash is regarded to be a key of potential colonists. In the vicinity of Ann Arbor, community element: green ash (n = 55), white ash (n Michigan – where EAB was first documented – the = 43), black ash (n = 14), and Oregon ash, Fraxinus ecological vacuum created by the loss of green ash latifolia Benth. (n = 12). Likewise, only eight species has been filled by spicebush (Lindera benzoin L.), of Fraxinus grow as the (lead) dominant tree species pawpaw (Asimina triloba Dunal), and prickly ash in 51 community types: green ash (n = 18 community (Zanthoxylum americanum Mill.). In the same area, Groups), Oregon ash (n = 10), white ash (n = 8), black one wetland formerly dominated by black ash has ash (n = 6), Carolina ash, F. caroliniana Miller (n = 3), changed into a monoculture of sedge (Carex L.). blue ash, F. quadrangulata Michx. (n = 3), singleleaf While both ash habitats changed structurally, neither ash F. anomala (n=2) and pumpkin ash (n = 1). gave way to invasive species (Anton Reznicek, Ecological contributions of each of North America’s pers. comm). In Indiana and Ohio, silky (Cornus 16 native Fraxinus, as identified in the U.S. National amomum Mill.) and gray dogwoods (C. racemosa Vegetation Classification and NatureServe Explorer Lam.) have flourished in communities where ashes (2014), are summarized in Table 1. (mostly green and black) have been lost (Jim In Table 2 we list 16 North American forest Bissell, Mike Homoya, and Jim McCormac, all community types where a Fraxinus is the dominant pers. comm). Invasive shrubs that have increased

BIOLOGY AND CONTROL OF EMERALD ASH BORER 20 CHAPTER 2: ECOLOGICAL IMPACTS OF EMERALD ASH BORER in post-EAB woodlands and bottomlands in Indiana often carry lower herbivore loads (Tallamy and and Ohio include multiflora rose (Rosa multiflora Shropshire, 2009) and as a consequence would not Thunb.), honeysuckle (Lonicera morrowii A. Gray support as many insectivorous birds, , or and three other honeysuckles), and glossy buckthorn other wildlife. However, we are compelled to interject (Rhamnus alnifolia L’Hér), but not to the extent that here that we generally find ash to be relatively many feared. As noted above, even in southeastern herbivore free. Low insect/pest loading is one reason Michigan where EAB was first documented, it is still that ashes are often chosen for city plantings (see too early to know what the ecological consequences of species treatments in Elias [1987]). In Tallamy and ash decline will be. Over time, sedge meadows, reed Shropshire (2009)’s compilation of the canary grass (Phalaris arundinacea L.) bottomlands, feeding on 1385 plant genera grown in the Mid- and various shrublands that arise following ash Atlantic States, Fraxinus ranks sixteenth in richness. die-off will give way to forest communities, but the Green and white ash trees planted in cities, towns, and composition of these replacement forests remains to parks in the western United States show little evidence be seen. It is also too soon to disregard Fraxinus as an of herbivory (DLW pers. observation). Likewise, in eventual component of replacement forests as virtually European woodlands, ash trees have been documented all EAB-affected communities in the Midwestern to have low herbivore loads relative to many other United States have seedling and sapling ash recruiting genera of forest trees (Fischbacher et al., 1998). Among presently (Anton Reznicek pers. comm). If native and four tree genera surveyed near Basel, Switzerland introduced natural enemies (e.g., Duan et al., 2013; (Carpinus L. [hornbeams], Fagus L. [beech], Fraxinus also see Chapters 8 and 9) are able to reduce emerald L. [ash], and Quercus L. [oak]), ash yielded only about ash borer density, Fraxinus could regain some of its half the caterpillar frass observed on other surveyed former ecological importance. genera. Moreover, while invasive plants generally have Where significant physiognomic changes occur, depauperate herbivore loads, at least Lonicera and e.g., where a woodland is initially replaced by a Rhamnus produce abundant fruits that are exploited graminoid wetland or shrubland, the biota, species by a range of birds and other vertebrates. As with all interactions, hydrology (Slesak et al., 2014), light ecological change, some species will benefit, and others regimen, nutrient cycling, vertebrate food value, and will suffer. other core ecosystem characteristics will be altered. A final note, brought to our attention by Anton Beyond the obvious loss of ash-specialized herbivores, Reznicek, is that considerable demographic differences structural changes in affected woodlands may change exist among North American Fraxinus as to their a forest’s suitability as breeding habitat and cover age and size of first reproduction. Green ash begins for resident vertebrates and invertebrates (e.g., see flowering and fruiting as a young tree – at diameters discussion of mole salamanders [Ambystoma species] frequently ignored by EAB, and thus the species has in pumpkin ash swamps later in this chapter). On a the potential to persist as young trees, e.g., in open smaller spatial scale, all of the above applies to the riparian and floodplain communities. By contrast, forest gaps that form when stands or glades of ash are white and pumpkin ash fruit later, and typically young killed by EAB. Such canopy gaps are noted to cause trees will succumb to EAB infestation before they microclimate effects altering, among other biota, can bear seed. Hence these species’ fate and those of populations (Gandhi et al., 2014). These their dependent herbivore faunas are likely to differ gaps also open up forests for invasions by plant species substantially from those of green ash. normally limited by light availability (Herms and McCullough, 2014). NORTH AMERICA’S SIXTEEN FRAXINUS When native plants are replaced by exotic species, such as glossy buckthorn and exotic honeysuckles in Below we provide a synopsis addressing the range, Indiana and Ohio (see above), there can be cascading preferred habitat, ecological importance, known consequences to higher trophic levels. Exotic plants susceptibility to EAB, and herbivore specialists of

BIOLOGY AND CONTROL OF EMERALD ASH BORER 21

CHAPTER 2: ECOLOGICAL IMPACTS OF EMERALD ASH BORER Total Species Species Total

4 1 1 3 1 2 3 1 17 46 29 20 10 10

Lepidoptera Lepidoptera

1 5 1 2 2 1

Hymenoptera Hymenoptera Hemiptera 1 2 4 2 5 8 1 1 7

Diptera Diptera

Specialist Herbivores Specialist Herbivores Coleoptera Coleoptera

3 1 Acari Acari 1 2 4 2 111 6 1 9 62 3 4 5 15 6 1 11 1 1 1 1 1 5 Approx. RegionApprox.

West Southwest Southwest East West South Midwest/East Midwest/East East Midwest/East Southwest Texas Southeast Southeast Texas Texas Texas East Midwest/Northeast Midwest/Northeast Total Total

discussion. Diets of southern and discussion. Diets Guilds and Specificity of Arthropod Herbivores Feeding

1 3 6 5 5 2 9 2 1 12 43 55 14 Co-dominate Co-dominate 1 5 2 2 1

Group Dominant Dominant 3 3 8 35 3 6 6 8 Forest Community Community Forest 10 2 18 37 Common Name Common Name California Ash California Ash Ash Fragrant Singleleaf Ash Ash Singleleaf 2 1 Blue Ash Ash Blue Oregon Ash Ash Oregon ash Chihuahuan White Ash Pumpkin Ash Ash Pumpkin Ash Green 1 Ash Velvet 4 Mexican Ash Ash Mexican Carolina Ash Ash Carolina Texas Ash Ash Texas Ash Gregg’s Goodding’s Ash Ash Goodding’s various ashes ashes various Black Ash

Fraxinus dipetala dipetala Fraxinus cuspidata Fraxinus Fraxinus anomala anomala Fraxinus Fraxinus quadrangulata quadrangulata Fraxinus Fraxinus latifolia latifolia Fraxinus papillosa Fraxinus Fraxinus americana americana Fraxinus Fraxinus profunda profunda Fraxinus pennsylvanica Fraxinus Fraxinus velutina Fraxinus berlandieriana berlandieriana Fraxinus Fraxinus caroliniana Fraxinus albicans Fraxinus albicans Fraxinus greggii greggii Fraxinus Fraxinus gooddingii spp. Fraxinus Fraxinus nigra nigra Fraxinus Clade Species Dipetalae Melioides Native ash species found North ash species found with their USNVC/NatureServe of Mexico . Native importance fauna . and tally of specialist herbivore Table1 Pauciflorae Pauciflorae In-the-wild host ranges are poorly known for most of the arthropod species in this table. We suspect that virtually all of the taxa that appear to be specialized in this be specialized appear to that virtually all of the taxa suspect that We species in this table. of the arthropod most for poorly known are ranges In-the-wild host Footnote: undersampling—see due to above than indicated ranges host broader have in fact, table, especially poorly known. are taxa western NA Fraxinus Fraxinus

BIOLOGY AND CONTROL OF EMERALD ASH BORER 22 CHAPTER 2: ECOLOGICAL IMPACTS OF EMERALD ASH BORER

Division Dune & American Eastern North & Shrubland Coastal Grassland Grass­ American Eastern North & Shrubland Meadow land, Grass­ American Eastern North & Shrubland Meadow land, Cool American Eastern North Forest Temperate Cool American Eastern North Forest Temperate Cool American Eastern North Forest Temperate North Southeastern Forest Temperate Warm American Flooded & Swamp Vancouverian Forest Flooded & Swamp Vancouverian Forest Flooded & Swamp Vancouverian Forest Flooded American Eastern North Forest & Swamp Flooded American Eastern North Forest & Swamp North Southeastern FloodedAmerican & Swamp Forest North Southeastern FloodedAmerican & Swamp Forest Cool American Eastern North Forest Temperate Cool American Eastern North Forest Temperate VA VA VA TX KY KY OR NC tion* GA, NC GA, NC MD, VA VA MD, OR , WA AR?, LA PA?, WV PA?, OR, WA? OR, WA? Distribu­ AL?, FL, GA? MD, VA, WV VA, MD, NatureServe NatureServe Status Critically imperiled Critically imperiled Imperiled Imperiled Critically imper­ iled? Imperiled Imperiled Critically imperiled Imperiled Imperiled Critically imperiled Critically imperiled - Imperiled Imperiled? Imperiled? Imperiled Critically imper­ iled?

- Solidago Solidago

- - / Piptochaetium

Woodland Juniperus virginiana Forest Allium cernuum Polygonum tenue - tenue Polygonum Schizachyrium scoparium Rhus aromatica Rhus aromatica - -

Helianthus divaricatus - / Helianthus hirsutus - cerifera / Morella taeda Pinus - pensylvanica / Carex - nutans Sorghastrum ) - trichocarpa spectabilis / Rubus

Talinum teretifolium / Talinum Forestiera acuminata / Phanopyrum acuminata gymnocarpon) / Forestiera Muhlenbergia sobolifera / Muhlenbergia / Symphoricarpos orbiculatus / Frangula caroliniana Herbaceous Vegetation Herbaceous

Forest texana) - ( Quercus americana - Ulmus heterophylla Populus Carya ovata Carya glabra Carya glabra Wooded Herbaceous Vegetation Herbaceous Wooded ) Wooded Herbaceous Vegetation Herbaceous ) Wooded Woodland spp. Woodland spp. / Elymus gigantea Tidal Forest Tidal

Woodland Woodland Forest ssp. ssp. Ecological sp. Fraxinus by dominated Community - gerardii / Andropogon americana Fraxinus Pycnanthemum tenuifolium avenaceum - americana Fraxinus - americana Fraxinus ulmifolia / Ptelea virginiana - Ostrya barbatum / Acer rubra - Ulmus nigra - Juglans americana Fraxinus trifoliata Forest dioica - Urtica deweyana / Carex latifolia Fraxinus Forest patens / Juncus latifolia Fraxinus ssp. balsamifera - Populus latifolia Fraxinus Forest alnifolia / Rhamnus balsamea - Abies nigra Fraxinus - ( Ulmus americana pennsylvanica Fraxinus silicicola var. - ( Carya aquatica pennsylvanica Fraxinus Forest Depression - pennsylvanica Fraxinus - Lithosper­ scoparium / Schizachyrium virginiana var. virginiana - Juniperus quadrangulata Fraxinus canescensmum / Arundinaria muehlenbergii - Quercus macrocarpa - Quercus quadrangulata Fraxinus gigantea - Juniperus americana virginiana Fraxinus - americana Fraxinus Opuntia humifusa / Physocarpus americana opulifolius Fraxinus dubia ( Phacelia associations. Virginia; WA - Washington; WV - West Virginia Virginia WV - West - Washington; WA Virginia; . See NatureServe Explorer (2014) for details on distribution, ranking, and additional plant details on distribution, and additional plant ranking, . See (2014) for NatureServe Explorer Fraxinus by dominated communities 2. Sixteen globally imperiled forest Table AL - Alabama, AR - Arkansas; FL - ; GA - Georgia; KY - Kentucky; LA - Louisiana; MD - Maryland; NC - North Carolina; OR - Oregon; PA - Pennsylvania; TX - Texas; VA - VA TX - Texas; - Pennsylvania; PA OR - Oregon; LA - Louisiana; MD Maryland; NC - North Carolina; KY - Kentucky; FL - Florida; GA Georgia; * AL - Alabama, AR Arkansas;

BIOLOGY AND CONTROL OF EMERALD ASH BORER 23 CHAPTER 2: ECOLOGICAL IMPACTS OF EMERALD ASH BORER

each North American Fraxinus. We group ash species hundreds of hectares, principally in the North by four phylogenetic sections based on Hinsinger American Great Lakes region (Fig. 5). Red maple- et al. (2013), beginning with the nominate Section black ash swamps cover thousands of hectares in Fraxinus, which is monotypic in the Americas and southern Canada and the northern United States. represented only by black ash (F. nigra). Of these Where Fraxinus nigra grows in monocultures or as four phylogenetic groupings, Section Melioides a dominant in either the canopy or subcanopy, the (in its broad sense) contains the most ecologically emerald ash borer represents a special threat. Black important, widespread species of ash; it is also the ash is considered to be the most vulnerable North most speciose section, with 10 North American American ash species as its range lies within the species. For the purposes of our evaluation, we make presumed climate niche for the beetle (Klooster et al., the assumption that all 16 North American species 2014; Liang and Fei, 2014; Leah Bauer pers. comm.). are vulnerable to the emerald ash borer, given initial Of additional concern is how plant community host preference studies and the uncertainty in climate composition will change in black ash stands post-EAB niche models for the insect (Sobek-Swant et al., 2012; invasion. Where the canopies are lost there is added Liang and Fei, 2014). However, it seems unlikely to risk that the local biota, forest structure, hydrology, us that the beetle will pose a major threat to the nine and other core ecological attributes will be harmed. species of western arid land ash (F. albicans Buckley, We recorded ten ash specialist herbivores from F. anomala Torr. ex S. Watson, F. berlandieriana black ash: one aphid, one seed , one sawfly, DC., F. cuspidata Torr., F. dipetala Hook. & Arn., F. one gracillariid leafminer, two noctuids, one pyralid, gooddingii Little, F. greggii, F. papillosa Lingelsh, and and three sphingids (Table 4). All but one of these F. velutina Torr.), given that EAB does not inhabit arthropods are also known from Section Melioides xeric and desert areas in its native range. sensu stricto Fraxinus, and especially F. americana, F. pennsylvanica, or both. The rarely encountered Section Fraxinus Canadian sphinx (Sphinx canadensis) () Black ash. Fraxinus nigra, a small tree that is the warrants special consideration. Tuttle (2007) wrote: most northern member of its genus, is distributed “In northeastern Indiana, [Sphinx canadensis] is across southern Canada from eastern Manitoba to closely associated with the understory of dense Newfoundland, south in mountains through West hardwood stands along the perimeter of wetlands. Virginia, in the Ohio River Valley to southern Indiana Over several seasons larvae were found exclusively on and Illinois, and in much of Iowa (Elias, 1987). Black black ash (Fraxinus nigra: Oleaeceae), although two ash is a major component of 14 forest community additional ash species [F. americana, F. pennsylvanica] Groups (NatureServe Explorer, 2014): six as a were present. Just as significant, larvae were never dominant tree species and eight as a co-dominate. It found on black ashes growing in open areas, is the dominant tree in one critically imperiled (G1) although those same trees supported the larvae of forest community type, the Fraxinus nigra - Abies two other sphingid species.” The northern range of balsamea /Rhamnus alnifolia forests of West Virginia the Canadian sphinx closely follows that of black ash. and Pennsylvania – two states with established EAB Southward the ’s range extends beyond that of populations. F. nigra, south of the Ohio River, where the species is Black ash grows in damp woods, bottomlands, rare and its Fraxinus hosts are unknown. swamps, and other wetlands; it tolerates considerable Section Pauciflorae inundation (Fig. 8). Common associates include black spruce (Picea mariana [Mill.] Britton, Sterns Goodding’s ash. Fraxinus gooddingii is a narrowly & Poggenb.), white cedar (Thuja occidentalis L.), distributed shrubby ash limited to southeastern tamarack (Larix laricina [Du Roi] K. Koch), birch Arizona and northern Sonora. It grows on rocky (Betula L.), and especially red maple (Acer rubrum slopes (often on limestone), in desert scrub, oak L.). It sometimes grows in nearly pure stands over woodlands, and riparian associations, scattered

BIOLOGY AND CONTROL OF EMERALD ASH BORER 24 CHAPTER 2: ECOLOGICAL IMPACTS OF EMERALD ASH BORER among other woody shrubs on lower canyon slopes; commonly planted as shade trees in yards, parks, and most records are from arroyos and canyon bottoms along streets (Elias, 1987; Burns and Honkala, 1990). from 1100 to 1500 meters (SEINet, 2014). EAB is thought to prefer other Fraxinus species over No previous literature has identified specialist white ash (Anulewicz et al., 2008), but F. americana is herbivores on this host. Jim Verrier and DLW still attacked and generally suffers very high mortality. found Philtraea elegantaria (Geometridae) feeding However, apparent resistance of white ash to EAB has on Goodding’s ash in Rock Corral Canyon in the recently been seen in Ohio, with some white ash trees Tumacácori Range. Sphinx libocedrus (Sphingidae) surviving in forests where green ash have succumbed feeds on this ash in Arizona (Tuttle, 2007), but species to the beetle (Jim Bissell pers. comm.). of Forestiera are presumed to be its principal hosts. White ash has the richest fauna of ash-specialized Gregg’s ash. Fraxinus greggii is found in the herbivores in North America – three times as many Trans-Pecos region of southwestern Texas south as any western ash. We list 46 species in Table 4, but at least to Hidalgo, Mexico. Gregg’s ash is often suspect that virtually all of the eastern specialists limited to riparian corridors in arroyos, canyon listed only from Fraxinus (without an associated bottoms, and along water courses from 400 to 1800 species epithet) probably use white ash. Specialist meters (Powell, 1998; SEINet, 2014). It grows on arthropods feeding on F. americana include 2 , 1 cliffs, rocky slopes, and canyon bottoms, frequently leaf beetle, 5 bark , 4 seed , 1 scarabaeid, on limestone. Throughout its range, it grows as a 1 agromyzid, 5 gall midges, 2 aphids, 6 mirids, 1 lace subdominant: typically as scattered plants, in desert bug, 3 sawflies, 2 inchworms, 2 gracillariid miners, 3 canyons and foothills, usually upslope from water. owlets (), 1 pyralid, 1 clearwing borer, and Noel McFarland (pers. comm.) found caterpillars 5 sphingids (Table 4). Sixteen herbivores are recorded of Philtraea paucimacula (Geometridae) in high only from white ash: 2 Hylesinus (), densities on Gregg’s ash near Laredo, Texas. Sphinx 2 Lignyodes (Curculionidae), 1 libocedrus (Sphingidae) is known from this ash, as (), 4 Dasineura (Cecidomyiidae), 1 well as Forestiera (Table 4). Prociphilus (Aphididae), 3 Tropidosteptes (), 1 Tethida (Tenthredinidae), 1 Hydrelia (Geometridae), Section Melioides and 1 Copivaleria (Noctuidae), although none of these are known to be strictly monophagous. White ash. Fraxinus americana is the second Green ash. Fraxinus pennsylvanica is a small­ most abundant and widespread native North to-medium-sized, fast growing tree that is the most American ash. It is a common component in many widely distributed ash species in North America, forest types, including bottomlands, open upslope extending from southwestern Saskatchewan to Cape woodlands (Fig. 11), and mixed hardwood forests. Breton Island, south to northern Florida and eastern Typically, white ash occurs in sites with enriched, but Texas. It is an abundant, ecologically important well drained, moist soils, where it grows in association species throughout much of this range and thrives with sugar maple (Acer saccharum Marshall), tulip in floodplains and other bottomlands. Elias (1987) tree (Liriodendron tulipifera L.), cherry (Prunus L.), notes that F. pennsylvanica is especially abundant beech, sweet gum (Liquidambar styraciflua L.), red through the Mississippi Valley. Green ash is the most maple, and willow (Salix L.). NatureServe Explorer common Fraxinus to grow as a dominant tree in the (2014) identifies eight ecosystems with white ash as a U.S. National Vegetation Classification (USNVC) dominant tree and 35 where it is a co-dominant. Of system. Fifty-five ecological community Groups the eight forest community Groups where white ash contain F. pennsylvanica as a dominant (n = 18) or co- is dominant, three are critically imperiled (G1) and dominant tree (n = 37) (NatureServe Explorer, 2014). four are imperiled (G2) (NatureServe Explorer, 2014) Green ash is the dominant tree in three forest Groups (Table 2). White ash seeds are eaten by many birds considered to be Critically Imperiled to Imperiled; and squirrels (Burns and Honkala, 1990) and it is all three of which are flooded swamp forests in the

BIOLOGY AND CONTROL OF EMERALD ASH BORER 25 CHAPTER 2: ECOLOGICAL IMPACTS OF EMERALD ASH BORER

southeastern United States (Table 2). In forests where Carolina ash. Fraxinus caroliniana is a small green ash is dominant, Burr and McCullough (2014) tree of the Atlantic Coast Plain from Virginia, south note that green ash is “unlikely to persist as a dominant through much of Florida, west to east Texas and species” after EAB invasion. Its seeds are an important southern Arkansas. It grows in swamps, wetlands, forage for a variety of birds, small mammals, and bottomlands, and other mesic to wet forest types. other wildlife; deer and moose (Alces alces L.) feed Like many other members of the genus, it does on new growth (Elias, 1987). Its strong wood is sold especially well in marl soils (SEINet, 2014). Carolina commercially for bats, oars, and tool handles but is not ash commonly grows with gums (Nyssa L.), hollies as desirable for wood working as white ash (Burns and (Ilex L.), sweet gum, sugarberry (Celtis laevigata Honkala, 1990). Willdenow), and others (Elias, 1987); it can be We record 29 Fraxinus-Oleaceae specialists from locally abundant as an understory member of bald green ash; thus, it ranks only behind white ash in cypress-tupelo swamps (SEINet, 2014). Fraxinus its importance to Fraxinus-specialist herbivores: 1 caroliniana is a dominant in nine forest community mite, 1 buprestid, 3 bark beetles (Hylesinus), 2 seed Groups: three as a dominant and six as a co-dominant weevils (Lignyodes), 1 agromyzid, 3 gall midges tree (NatureServe Explorer, 2014). It occurs in (Cecidomyiidae), 1 aphid, 5 mirids, 1 sawfly, 4 dense monospecific stands in coastal plain areas gracillariid miners, 2 owlets (Noctuidae), 1 pyralid, of Louisiana and Texas (in “Fraxinus caroliniana 1 clearwing borer (Sesiidae), and 3 sphingids. Seasonally Flooded Forests”), which are deemed Seven species are recorded only from green ash: G2G3 imperiled-vulnerable communities. Only one one undescribed Dasineura (Cecidomyiidae), two herbivore, the sphingid moth Ceratomia undulosa, Tropidosteptes (Miridae), and the two gracillariids an oligophage on Oleaceae, is recorded from this Marmara basidendroca and Marmara corticola. The last Fraxinus. The lack of feeding records for this ash two of these, so far as known, are monophages, at least is almost certainly an artifact of limited sampling, at the type locality and surrounding areas of upstate presumably because its preferred habitat is in swamps (Fitzgerald, 1973; Terry Fitzgerald pers. and wetlands and it is not a commercially important comm.). See also Imperilment Risk Rating discussion. ash. Mexican ash. Fraxinus berlandieriana is a small Fragrant ash. Fraxinus cuspidata is a shrub tree that occurs from the vicinity of Austin, Texas (or sometimes small tree) that is found in scattered southward and westward into Mexico. The core of populations across the southwestern United States its range is in the Mexican states of Coahuila, Nuevo from northwestern Arizona to western Texas and Leon, and Tamaulipas. It grows in moist canyons and south into Mexico, mostly in the foothills of desert along streams and rivers of the Rio Grande Plains ranges (SEINet, 2014). It prefers well-drained, rocky and southern prairies (Vines, 1984) and is frequently soils in canyons, and north facing cliffs of limestone, planted as a shade tree in parks and cities. Two sandstone, or igneous soils (Elias, 1987; Powell, 1998; community types in Texas include Mexican ash as a co- SEINet, 2014). It tends to occur in low densities dominate (n = 2), one of which is considered critically intermixed among oaks, leguminous trees, and other imperiled/imperiled (G1G2) (NatureServe Explorer, woody plants. 2014). Two specialist herbivores have been recorded from this small ash: DLW collected caterpillars of Ash specialists reported from the United States a Sympistis heterogena (Noctuidae) feeding on new include the buprestid Trigonogya reticulaticollis, the spring leaves in May 2014 and a second unidentified chrysomelid Capraita sexmaculata, and the seed weevil noctuid2 species in July of 2014. Lignyodes helvolus. While Mexican ash is the only Oregon ash. Fraxinus latifolia is a Pacific reported host for T. reticulaticollis, based on the biology Coast tree that grows from the Olympia area in of related metallic wood boring beetles, it is probable that its host range will be found to include other 2 The caterpillar, seen in late July, was banded with smoky red in Oleaceae. the early instars and is a new foliage specialist.

BIOLOGY AND CONTROL OF EMERALD ASH BORER 26 CHAPTER 2: ECOLOGICAL IMPACTS OF EMERALD ASH BORER western Washington south to the San Francisco on north-facing cliffs (SEINet, 2014). Like other Bay Area (California) and southern Sierra Nevada, desert ashes, it occurs as scattered plants in woody as well as in disjunct populations in southern associations near water or in microhabitats with California’s mountains. The tree grows in “moist reduced moisture stress. No ash specialists are rich soils along streams and rivers, and in canyons” currently recorded on F. papillosa. In Hinsinger et to 500 meters (Elias, 1987). It is an abundant and al.’s (2013) study, Chihuahuan ash grouped with ecologically important tree along riparian corridors Oregon ash (F. latifolia), in a clade outside of all other throughout much of its range, and is the only members of the Melioides sensu stricto section of the commercially important Fraxinus in western North genus. Because of this phylogenetic difference and America. Oregon ash is a dominant tree in ten forest its taxonomic and geographic isolation, Chihuahuan community Groups and a co-dominant in two others. ash strikes us as a likely candidate for hosting unique While it will grow in monocultures, at least along herbivores. streams and in floodplains, it more typically grows in Pumpkin ash. Fraxinus profunda grows in mixed hardwood stands with maple, alder (Alnus scattered locations across eastern North America, with Mill.), poplar (Populus L.), and willow; Elias (1987) most occurrences along the Atlantic Coastal Plain and also adds California laurel (Umbellularia californica through the Ohio and Mississippi River Valleys. It is [Hook. & Arn.] Nutt.) and grand fir (Abies grandis restricted to very wet soils that include bottomlands, [Douglas ex D. Don] Lindley) as associates of F. floodplains, swamps (especially bald cypress and tupelo latifolia. Three forest communities in the Pacific swamps), and coastal marshes (Elias, 1987; Burns and Northwest dominated by Oregon ash are regarded Honkala, 1990; Nesom, 2010) (Figs. 7, 9). Fraxinus as critically imperiled/imperiled in NatureServe profunda is a dominant (n = 1) and co-dominant (n = Explorer (2014) (See Table 2). 4) in five eastern forest community types, one of which We record 17 species of ash specialists from is imperiled (NatureServe Explorer, 2014). This ash Oregon ash, many for the first time: 2 mites, 2 species is quite susceptible to EAB; devastated would bark beetles, 2 seed weevils, 1 aphid, 2 mirids, 1 not be an overstated descriptor for some local Ohio tingid, 2 sawflies, and 1 inchworm (Geometridae), preserves where pumpkin ash once grew as a dominant 1 gracillariid leafminer, 1 owlet moth, and 1 ermine species (Jim Bissell pers. comm.) moth (Yponomeutidae). Five of these are only known Only one arthropod species is recorded from F. from Oregon ash: Hylesinus oregonus (Curculionidae: profunda: a mite that is not a specialist on pumpkin Scolytinae); Lignyodes auratus (Curculionidae: ash (Table 1). No doubt pumpkin ash’s poorly sampled ); Philtraea latifoliae (Geometridae), herbivore fauna is a reflection of the difficulty involved Caloptilia n. sp. (Gracillariidae), and Sympistis fortis with sampling the inundated wetlands where this (Noctuidae). We suspect that their strict monophagy tree thrives. Certainly it is a candidate for more study, is either an artifact of undersampling or because especially given its susceptibility and because its range Fraxinus specialists whose ranges are restricted falls entirely within the predicted climatic niche of to the Sierra and areas north of San Francisco are EAB. monophagous simply because F. latifolia is the only Velvet ash. Fraxinus velutina is a small ash of the ash that grows where they occur. southwestern United States and northwestern Mexico. Chihuahua ash. Fraxinus papillosa is a Its U. S. range is concentrated in Arizona and extends geographically restricted ash that grows to become a westward through arid portions of southern California, small tree. It is found in small populations through Nevada, , and eastward into the Trans-Pecos desert ranges of southeast Arizona, southwestern region of Texas. It grows near streams, rivers, washes, New Mexico, the Chinati Mountains of west Texas, and other areas of reduced moisture stress. Common and southward into the Sierra Madre Occidental associates include hackberry (Celtis occidentalis L.), Mountains of Mexico (Elias, 1987; Powell, 1998). oak, poplar, sycamore (Platanus L.), and willow. It Typically F. papillosa grows in canyon bottoms and reaches greatest importance in mesic stretches of

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canyons between 1200 and 1600 meters where it “can host for Philatraea utahensis Buckett. form almost pure stands” (Elias, 1987). Velvet ash is California or two-petal ash. Fraxinus dipetala listed as a co-dominant tree in five forest community is a shrubby ash that grows in scattered populations Groups in NatureServe Explorer (2014). in central and southern California’s coastal ranges We record 20 ash specialist herbivores from and the foothills of the Sierra Nevada Mountains. It velvet ash, the third most of any Fraxinus species thrives either on slightly acidic granite slopes, often and the greatest number for any western species: 2 near watercourses, or on slightly alkaline clays in mites, 1 bark beetle, 2 seed weevils, 2 scarab beetles, chaparral associations. Below 500 meters, it is usually 2 aphids, 2 scale , 2 plant bugs, 1 stink big, 1 restricted to the upslope vicinities of washes and lace bug, 2 owlet (Noctuidae), and 3 sphingids. watercourses. Above this, California ash occurs in Most of these are known only from F. velutina: the chaparral and other foothill communities, typically as mite Brevipalpus cardinalis (Tenuipalpidae), the a scattered element growing with Pinus L. (pine), oak, weevil Lignyodes arizonicus (Curculionidae), the Arctostaphylos Adans. (manzanita), and others (Elias, beetles granti and Xyloryctes thestalus (both 1987; SEINet, 2014). Fraxinus dipetala is listed as a Scarabaeidae), the armored scales Diaspidiotus fraxini co-dominant (along with Prunus ilicifolia [Nutt. ex and Diaspis fraxini (both Diaspididae), the plant Hook. & Arn.] Walp.) in one U.S. forest community bug Tropidosteptes illitus (Miridae), the stink bug B. Group by NatureServe Explorer (2014). rolstoni (Pentatomidae), and the moths Chloronycta The herbivore fauna of California ash is not well tybo and Sympistis punctilinea (both Noctuidae) known – the plant is easily overlooked, and rarely and Ceratomia sonorensis and Sphinx near chersis common. Four ash-specialist herbivores are recorded (both Sphingidae). In the Sonoran and Chihuahuan from California ash: 1 mite, 1 aphid, 1 inchworm deserts of the western United States, velvet ash is (Geometridae), and 1 ermine moth (Yponomeutidae). the only Fraxinus considered to be an ecologically Two moths are known exclusively from this ash: dominant or, more commonly, co-dominant species. Philtraea surcaliforniae (Geometridae) and Zelleria Consequently, the loss of velvet ash would have an near semitincta (Yponomeutidae) (Buckett, 1970) impact on all southwestern Fraxinus herbivores, given (Table 4). that it accounts for most of the ash biomass in the Blue ash. Fraxinus quadrangulata is a small to southwestern United States. large tree (Fig. 6) whose core distribution is in the Midwestern United States from eastern Kansas to Section Dipetalae extreme southern Ontario, south into Arkansas Singleleaf ash. Fraxinus anomala is a shrub or and northern Alabama. Blue ash grows in upland small spreading tree that grows in canyons, desert woods and on slopes usually over limestone, drainages, washes, and along water bodies of the but also in moist woodlands and evidently even North American Great Basin region. It exists as bottomland forests (Elias, 1987). Typically it occurs scattered stands in western Colorado, Utah (where it as a subdominant species (n = 3) intermixed in is widespread), southern Nevada, and southeastern stands dominated by oak, mockernut hickory (Carya California, central and northern Arizona, and tomentosa Sarg.), sweet gum, and others. It is listed extreme northwestern New Mexico (SEINet, 2014). as the dominant tree species in three Kentucky forest Fraxinus anomala is a common riparian element community types (NatureServe Explorer, 2014): between 600-1,900 meters. NatureServe Explorer critically imperiled (G1) blue ash-oak savannah (2014) recognizes singleleaf ash as a dominant (n woodlands (Bryant et al., 1980; NatureServe Explorer, = 2) or co-dominant (n = 1) in three southwestern 2014); imperiled (G2) blue ash - eastern red-cedar plant community associations (all three are currently / little bluestem - hoary puccoon woodlands; unranked or unrankable due to lack of data). We do and possibly no longer extant (GH) Fraxinus not know of any specialist ash herbivores from F. quadrangulata - Quercus macrocarpa / Arundinaria anomala, although we suspect that it is the primary gigantea ssp. gigantea wooded shrublands. Fraxinus

BIOLOGY AND CONTROL OF EMERALD ASH BORER 28 CHAPTER 2: ECOLOGICAL IMPACTS OF EMERALD ASH BORER quadrangulata is believed to be the most EAB- EFFECTS ON VERTEBRATES resistant Fraxinus in eastern North America and, consequently, its relative abundance may increase Martin et al. (1951) regarded North American ashes in woodlands and forests where other ash succumb to be “only of moderate importance to wildlife.” to the beetle (Anulewicz et al., 2007; Tanis and No vertebrates are monophagous or otherwise McCullough, 2012). exclusively dependent on ash or ash-dominated Blue ash has a surprisingly modest herbivore communities, although some local populations might fauna. Only one ash specialist is recorded from this be energetically or ecologically dependent on ash ash, the moth Plagodis kuetzingi (Geometridae), simply due to the fact that Fraxinus are the dominant which also occurs on F. americana and presumably or co-dominant trees in the woodland or forest types other ashes. Robinson et al. (2014) lists only two where they live (northern black ash swamps provide generalist Lepidoptera for blue ash. Its diminished compelling examples of such) (Fig. 5). A summary herbivore fauna no doubt is a reflection of its of vertebrates using ash as a food source is given in taxonomic isolation: it is the only member of Table 3, with the proviso that the published literature the Dipetalae Section in eastern North America is greatly biased towards game species. We also note (Hinsinger et al., 2013). Given the above, perhaps it is that the literature for vertebrates is largely qualitative, not surprising that blue ash is more resistant to EAB with little data on the importance of ash relative to than other eastern members of the genus. Given that other food resources; we suspect that the paucity of it represents a unique taxonomic entity in eastern studies addressing the ecological importance of ash North America, blue ash is a candidate for a focused may be a reflection of the fact that, by itself, ash trees herbivore survey, especially for cecidomyiids, leaf or rarely serve as a core food or sheltering resource. bast miners, and other herbivores known to show We know most about the six species of Fraxinus extreme monophagy. where ash has been identified as being a community Texas ash. Fraxinus albicans (= F. texensis) is dominant tree species: black, blue, green, Oregon, a small drought-tolerant relative of white ash that pumpkin, and white. Among mammals, ungulates grows on limestone from the Arbuckle Mountains of and lagomorphs (rabbits) are often mentioned as southern Oklahoma southward across the Edwards feeding on non-woody tissues of ash. Northward, Plateau into south central Texas (Vines, 1984; Elias, green and especially black ash are browse sources 1987). It is a co-dominant (with oak) in two forest for moose (Elias, 1987) and the white-tailed deer community types (NatureServe Explorer, 2014). We (Odocoileus virginianus Zimmermann). A wide know of only two reports of specialist herbivores range of rodents, but especially chipmunks, mice, from Texas ash: 1 mite and 1 weevil, neither of which and squirrels, consume the winged seeds. And, while is monophagous on F. albicans. Robinson et al. (2014) we found little mention of such in the literature, does not list any lepidopterans from Texas ash. undoubtedly North American beavers (Castor The dearth of feeding records for this ash is surely canadensis Kuhl) consume ash across their range. a reflection of its limited geographic distribution Ash seeds are regarded to be of moderate importance and thus a sampling artifact. Presumably because to seed-eating, woodland or forest dwelling birds of its close relation to F. americana (Vines, 1984), such as cardinals, chickadees, finches, and grosbeaks. F. caroliniana, and other members of the Melioides Grouse, quail, and turkey are ground feeders sensu stricto section of the genus (Hinsinger et known to consume ash seeds (Martin et al., 1951). al., 2013), F. albicans will be found to share many Given the propensity of the genus for wetlands, the herbivores with other Fraxinus. importance of ash seeds to waterfowl has likely been underestimated – we only found specific mention of wood ducks (Aix sponsa [L.]) feeding on ash samaras (Martin et al., 1951). In forests where Fraxinus are dominants, ashes also provide cover (e.g., for

BIOLOGY AND CONTROL OF EMERALD ASH BORER 29 CHAPTER 2: ECOLOGICAL IMPACTS OF EMERALD ASH BORER large seed crops seed crops large wildlife” to value “little Comments Comments Elias, 1987; Burns and Honkala, 1990 Honkala, and Elias, 1987; Burns 1990 Honkala, and Elias, 1987; Burns Elias, 1987 Elias, 1987; Burns and Honkala, 1990 Honkala, and Elias, 1987; Burns Elias, 1987 2005 Block, 1951; Rhoads et al., and Martin Elias, 1987; Burns and Honkala, 1990 Honkala, and Elias, 1987; Burns Literature 1990 Honkala, and Elias, 1987; Burns

wood ducks, quail, turkey, cardinals, finches finches cardinals, turkey, quail, wood ducks, birds song turkey, grouse, wood ducks, wood ducks, many other birds birds other many wood ducks, songbirds turkey, quail, wood ducks, many birds birds many birds song turkey, grouse, wood ducks, Birds Birds bobwhite, grouse, turkey, quail, wood ducks, grosbeaks purple finch), (esp. (esp. finches songbirds other grosbeak), cardinals, pine

white tail deer, mule deer, deer, mule tail deer, white squirrels, moose, rabbits, rodents other and moose, tail deer, white small mammals white tail deer white tail deer white chipmunks white tail deer, moose, tail deer, white small mammals deer squir­ ground rabbits, deer, rodents other rels, Mammals Mammals squirrels fox tail deer, white Fraxinus velutina Fraxinus pennsylvanica pennsylvanica Fraxinus nigra Fraxinus Fraxinus papillosa Fraxinus profunda Fraxinus quadrangulata Fraxinus spp. Fraxinus Fraxinus nigra nigra Fraxinus Fraxinus anomala Fraxinus berlandieriana Fraxinus Fraxinus caroliniana cuspidata Fraxinus dipetala Fraxinus Fraxinus gooddingii greggii Fraxinus Ash Tree Fraxinus albicans americana Fraxinus . Fraxinus on feed that 3. Vertebrates Table

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13 14 15

16 17 18

19 20

Figures 13 - 20. Ash-specialist insects that represent some of the larger taxa threatened by EAB. (13) Tropidosteptes cardinalis, Family Miridae (Photo courtesy Mike Quinn); (14) Leptoypha mutica, Family . (Photo courtesy Tom Murray); (15) Philtraea elegantaria, Family Geometridae (Photo DLW); (16) Sphinx franckii, Family Sphingidae (Photo DLW); (17) Sphinx kalmiae, Family Sphingidae (Photo DLW); (18) Dynastes granti, Family Scarabaeidae (Photo courtesy Margarethe Brummerman); (19) Lignyodes helvolus, Family Curculionidae (Photo courtesy Bill Johnson); (20) Xylorctes thestalus, Family Scrarabeidae. (Photo courtesy Richard Hayes)

BIOLOGY AND CONTROL OF EMERALD ASH BORER 31 CHAPTER 2: ECOLOGICAL IMPACTS OF EMERALD ASH BORER

moose [Gould and Bauer, 2009]) and above-ground species may also allow fruit-producing shrubs such structure important to vertebrate wildlife for roosting as Cornus, Lonicera, and Rhamnus (two of which are, and nesting. Ash trees with broken tops or otherwise unfortunately, non-native invasives) to colonize and damaged stems sometimes form cavities which are thrive, the fruits of which will benefit a variety of fruit- used by a variety of birds, bats, and other . eating birds and mammals. One ash community type warrants special mention, i.e., woodland pools dominated by pumpkin EFFECTS ON ASH-FEEDING ash (F. profunda) (Figs. 7, 9). In southeastern Michigan, INVERTEBRATES portions of Ohio, and presumably elsewhere, pumpkin ash is sometimes the sole or dominant tree that We identify 98 Fraxinus-dependent invertebrate her­ grows in the standing water of vernal and permanent bivores (or inquilines) as potentially threatened by the (fishless) forest pools and as such provides all or much spread of EAB, 45 of which are reported here for the first of the canopy closure. Ash-dominated woodland and time (Figs. 13-20). Because our compilation of Fraxinus forest pools are an important breeding habitat for feeders was a bottom-up tabulation for all insects and five species of mole salamanders in the Midwestern mites, built upon the collective knowledge of more than United States (spotted [Ambystoma maculatum Shaw], 80 taxonomic experts, we feel the data in Table 4 offer Jefferson [A. jeffersonianum (Green)], blue-spotted a unique look at the taxonomic distribution of ash-spe­ [A. laterale Hallowell], small-mouthed [A. texanum cialist herbivores from the estimated 70,000 species Matthes], and marbled [A. opacum Gravenhorst]), of North American insects (Arnett, 2000) and Acari and an occasional breeding habitat for the eastern (mites). Specialist herbivores that would be imperiled or tiger salamander (A. tigrinum Green). Many of these extirpated in the United States and Canada by the loss salamanders are important conservation targets of Fraxinus include mites (n = 6) and members of five that receive legal protection. Of these, the marbled insect orders: Lepidoptera (n = 32), Hemiptera (n = 25), salamander may be the most vulnerable because Coleoptera (n = 24), Diptera (n = 9), and Hymenoptera females lay their in mud during the fall, in (n = 3) (Fig. 21). The most speciose lineage of metazoans anticipation of autumn and winter rains (Degraaf and on the planet, beetles, had fewer specialists than Lepi­ Rudis, 1986; Pfingsten et al., 2013; Kenney, 1995). doptera and essentially equivalent richness to that of He­ Greater sun exposure due to canopy loss could affect miptera. While the focal taxon of our study was Fraxinus the hydrology of wetlands, especially in the fall, and related Oleaceae, we suspect that the proportions before rains and snow melt have had a chance to fill represented here are likely to apply across most temper­ them. Most worrisome is that, in Ohio, pumpkin ash ate woody, broadleaf plant taxa, and may well apply to is sometimes the only tree found growing in these other continental biogeographic provinces as well. saturated to flooded soils where mole salamanders and Four genera contain six or more species that will be other wildlife breed (Jim McCormac, pers. comm.). threatened by the spread of EAB; in decreasing diversity While much wildlife will experience detrimental these include Tropidosteptes plant bugs (Miridae) (n = effects from ash elimination, no doubt some 14) (Fig. 13), Hylesinus bark beetles (Curculionidae) (n = vertebrates will benefit. Where EAB has invaded in 7), Lignyodes seed weevils (Curculionidae) (n = 7) (Fig. the Midwestern United States, woodpecker numbers 19), and Sphinx hawkmoths (n = 6) (Sphingidae) (Figs. have spiked in the vicinity of infestations because 16, 17). Four other genera contain noteworthy radiations EAB larvae serve as an abundant, easily harvested on Fraxinus: Dasineura gall midges (Cecidomyiidae), food resource and dying ash provide nesting trees Prociphilus aphids (Aphididae), Philtraea inchworms for the birds (Cappaert et al., 2005b; Koenig et (Geometridae) (Fig. 15), and Sympistis sallows al., 2013). Similarly, standing dead ash trees are (Noctuidae). preferred roosting and nesting sites for many other vertebrates and will, for a limited time, benefit wildlife posthumously. Canopy gaps left by these dying ash

BIOLOGY AND CONTROL OF EMERALD ASH BORER 32 CHAPTER 2: ECOLOGICAL IMPACTS OF EMERALD ASH BORER

Figure 21. At risk arthropod herbivores by taxon and imperilment score.

Charismatic Arthropod Herbivores low or moderate endangerment risk in their compilation are treated here as imperiled The sphingids or hawkmoths deserve special mention ash specialists: Ceratomia undulosa, Manduca because of their beauty, size, and popularity (with jasminearum, Sphinx canadensis, S. chersis, S. franckii moth watchers, photographers, and collectors), (Fig. 16), and S. kalmiae (Fig. 17). We add three as well as their vulnerability and importance in resident hawkmoths not mentioned vertebrate diets. Furthermore, the degree of sphingid in previous ecological assessments of the North imperilment has heretofore been underestimated. American sphingid fauna: Ceratomia sonorensis, Gandhi and Herms (2010) listed eight native and one Sphinx libocedrus, and S. near chersis. We assign exotic species of sphingids as potentially affected by endangerment risks to these nine ash-dependent EAB invasion; the only sphingid that they ranked as as polyphagous taxa if one consults treatments such as Covell highly to moderately imperiled by EAB was Manduca (2005), Wagner (2005), Tuttle (2007), Robinson et al. (2014), etc. brontes, a Caribbean and South Florida species that But if records >50-years old or exceptional reports are excluded, feeds on Tecoma stans (L.) Juss. ex Kunth in the all are better regarded as imperiled Fraxinus or Oleaceae specialists. Older literature is fraught with misidentifications Bignoniaceae. Fraxinus host records for the moth and erroneous host records: For example, the host records of were determined to be erroneous by Robinson et al. Ulmus for S. franckii and M. jasminearum are almost certainly based on misidentifications of larval Ceratomia amyntor (Geyer, (2002); we exclude this species from our assessment. 1835), an elm feeder. To the best of our knowledge (and that of Six sphingids that were regarded as generalists3 of Jim Tuttle in litt.) none of these six species has been collected on host plants outside of the Oleaceae in our lifetimes, and all 3 Sphinx canadensis, S. chersis, S. kalmiae, and S. franckii, Man- but S. kalmiae (which can be found on other Oleaceae) are best duca jasminearum, and Ceratomia undulosa could be classified considered Fraxinus specialists.

BIOLOGY AND CONTROL OF EMERALD ASH BORER 33 CHAPTER 2: ECOLOGICAL IMPACTS OF EMERALD ASH BORER

(Richard (Richard , but that is is that , but Comments Comments Confirmed larval host; (1937) recorded Knull F. collected on adults caroliniana of F. a misidentification berlandieriana Host in litt.). Westcott study. needs more range Probably obligatory on on obligatory Probably diet includes adult ash; Oleaceae hosts. other Larvae feeders; root certainlyalmost spe­ cialists. Larvae feeders; root certainlyalmost spe­ cialists. Reference(s) Reference(s) MacRae, 2006; Ted 2006; Ted MacRae, MacRae unpubl. Richard data; in litt. Westcott Felt, 1918; Baker et Felt, Och­ 1996; Ron al., ua pers. comm. 1918; Baker et Felt, 1996 al., 1975; et al., Jeppson Baker 1996 et al., DeLeon, 1961; Suigong, Baker and Ochua 1988; Ron pers. comm. Banks, 1915; Baker 1988; Suigong, and 2007 Johnson, 1987; Ted Bright, MacRae unpubl. Richard data; in litt. Westcott Clark et al., 2004; et al., Clark in litt.; Clark Shawn Ed Riley in litt. Clark et al., 2004; et al., Clark in litt.; Clark Shawn Ed Riley in litt. Clark et al., 2004; et al., Clark in litt. Clark Shawn Blatchley and Leng, and Blatchley 1982; 1916; Wood, 2014, Atkinson, Cognato Anthony in litt. root root leaves leaves leaves leaves leaves Guild leaf gallleaf cambium cambium unknown flower gall flower recently dead wood recently High High High High High High High High High? High? Risk Rating Risk Rating High to Moderate Moderate to High Moderate to High

Confirmed Hosts ConfirmedHosts Fraxinus americana, americana, Fraxinus latifolia F. Fraxinus americana, americana, Fraxinus latifolia, F. pennsylvanica F. dipetala, Fraxinus velutina F. Fraxinus, F. profunda velutina, Fraxinus albicans F. , pennsylvanica Fraxinus Ligustrum berlandieriana Fraxinus Chionanthus virginicus, virginicus, Chionanthus F. americana, Fraxinus berlandieriana, ligustrinaForestiera Fraxinus Fraxinus Fraxinus Fraxinus Fraxinus americana, americana, Fraxinus pennsylvanica F.

(Schaeff er) (Schaeff

Scientifi c Name Name c Scientifi (Felt) (Felt) ora fraxinifl Aceria (Garman) (Garman) fraxini Aceria califraxini Tegolophus (Keifer) fraxini Brevipalpus DeLeon Brevipalpus cardinalis (Banks) Gory subcinctus Agrilus Trigonogya reticulaticollis Capraita sexmaculata sexmaculata Capraita (Illiger) Trichaltica tibialis Trichaltica (Jacoby) Trichaltica scabricula Trichaltica (Crotch) (Say) (Say) aculeatus Hylesinus Family Family Eriophyidae Eriophyidae Eriophyidae Eriophyidae Eriophyidae Tenuipalpidae Tenuipalpidae Buprestidae Chrysomelidae Chrysomelidae Chrysomelidae Curculionidae Curculionidae Taxon Acari Acari Acari Acari Acari Coleoptera Coleoptera Coleoptera Coleoptera Coleoptera Coleoptera Coleoptera Coleoptera Coleoptera Coleoptera and related Oleaceae. and related on Fraxinus 4. Specialist herbivores Table

BIOLOGY AND CONTROL OF EMERALD ASH BORER 34 CHAPTER 2: ECOLOGICAL IMPACTS OF EMERALD ASH BORER Comments Comments Reference(s) Reference(s) Wood, 1982; Langor 1982; Langor Wood, 1993; Hergert, and 2014; Atkinson, Cognato Anthony in litt. 2014; Atkinson, Cognato Anthony in litt. Leng, and Blatchley 1982; 1916; Wood, 2014; Atkinson, Cognato Anthony in litt. 2014; Atkinson, Cognato Anthony in litt. 1982; Burns Wood, 1990; Honkala, and 2014; Atkinson, Cognato Anthony in litt. 1982; Atkin­ Wood, 2014; Anthony son, in litt. Cognato 1980; RobertClark, pers. Anderson comm. 1980; RobertClark, pers. Anderson comm. Knull, 1932; Clark, 1932; Clark, Knull, 1980; Robert An­ pers. comm. derson 1980; RobertClark, pers. Anderson comm. Clark, 1980; RobertClark, pers. Anderson comm. seed seed seed seed seed Guild cambium cambium cambium cambium cambium cambium High High High High High High High Moderate Moderate Risk Rating Risk Rating High to Moderate Moderate to High Moderate to High High to Moderate Moderate to High

Confirmed Hosts ConfirmedHosts Fraxinus americana, americana, Fraxinus pennsylvanica F. americana Fraxinus albicans, Fraxinus Forestiera, Olea Ligustrum, latifolia Fraxinus americana Fraxinus Fraxinus velutina latifolia Fraxinus americana, Fraxinus nigra, F. pennsylvanica, F. Syringa Fraxinus americana, americana, Fraxinus latifolia, F. pennsylvanica, F. velutina, F. europaea Olea Fraxinus americana americana Fraxinus americana, Fraxinus berlandieriana, F. Syringa nigra, F.

Clark

Scientifi c Name Name c Scientifi Hylesinus criddlei Hylesinus ( Swaine) fasciatus Hylesinus LeConte mexicanus Hylesinus (Wood) oregonusHylesinus (Blackman) Hylesinus pruinosus Eichhoff arizonicus Lignyodes (Sleeper) auratus Lignyodes bischoffi Lignyodes (Blatchley) Hylesinus californicusHylesinus (Swaine) Lignyodes fraxini fraxini Lignyodes ( LeConte) helvolus Lignyodes (LeConte) Family Family Curculionidae Curculionidae Curculionidae Curculionidae Curculionidae Curculionidae Curculionidae Curculionidae Curculionidae Curculionidae Curculionidae Curculionidae Curculionidae Curculionidae Taxon Coleoptera Coleoptera Coleoptera Coleoptera Coleoptera Coleoptera Coleoptera Coleoptera Coleoptera Coleoptera Coleoptera Coleoptera Coleoptera Coleoptera (continued) 4. (continued) Table

BIOLOGY AND CONTROL OF EMERALD ASH BORER 35 CHAPTER 2: ECOLOGICAL IMPACTS OF EMERALD ASH BORER Comments Comments Larvae be general­ may feed on adults but ists absent or beetleash; rare (Bill Fraxinus from away in litt.). Warner Adults feed (and mate?) mate?) feed (and Adults into gnawed wounds at ash shoots. See in Rat- discussion e 2009. cliff See in Rat- discussion e 2009. cliff Phytophagous inquiline inquiline Phytophagous Dasineura in galls of tumidosae. Samara exocarp gall; Samara undescribed gregarious; evidently; Europe­ near species C. marchali. an Reference(s) Reference(s) Clark, 1980; RobertClark, pers. Anderson comm. 1980; RobertClark, pers. Anderson comm. 1980; RobertClark, pers. Anderson comm. Leng, and Blatchley et 1916; Arnett 2002; Charles al., in litt.; O’Brien Westcott Richard in litt. in litt.; Bill Warner pers. Sullivan Pat Margarethe comm.; pers. Brummerman comm. Glaser, 1976 Glaser, Ratcliff e, 2009 Ratcliff Ratcliff e, 2009 Ratcliff Solomon, 1995; Solomon, er in litt. Sheff Sonja 1989 Gagné, Gagné, 1989 Gagné, root root seed seed seed seed Guild cambium samara gall samara new shoot feeder new shoot new shoot feeder new shoot leaf gall (inquiline) gallleaf (inquiline) High High High High High High High High ments ments ments ments Risk Rating Risk Rating High-see com­ High-see com­ High to Moderate Moderate to High High to Moderate Moderate to High

and

, Syringa Confirmed Hosts ConfirmedHosts Fraxinus americana, F. F. americana, Fraxinus berlandieriana, Syringa nigra, F. Fraxinus latifolia, F. F. latifolia, Fraxinus , pennsylvanica velutina F. Fraxinus americana Fraxinus Fraxinus velutina Fraxinus Fraxinus Fraxinus americana americana Fraxinus Fraxinus velutina other likely in species, esp. Fraxinus Mexico Fraxinus americana, F. F. americana, Fraxinus pennsylvanica americana F. Fraxinus, Fraxinus pennsylvanica pennsylvanica Fraxinus Felt) Felt)

Bates

(Le-

n. sp. n. sp.

sp.

Scientifi c Name Name c Scientifi Conte) Lignyodes helvolus helvolus Lignyodes Lignyodes horridulus Lignyodes (Casey) ocularis Lignyodes (Casey) (Dietz) armatus Psomus politus)(including Horn granti Dynastes Dynastes tityus L. tityus Dynastes Xyloryctes jamaicensis jamaicensis Xyloryctes (Drury) Xyloryctes thestalus Phytobia ) fraxini (including Contarinia thalactri ( Contarinia Family Family Curculionidae Curculionidae Curculionidae Curculionidae Curculionidae Curculionidae Scarabaeidae Scarabaeidae Scarabaeidae Scarabaeidae Agromyzidae Agromyzidae Cecidomyiidae Cecidomyiidae Cecidomyiidae Cecidomyiidae Taxon Coleoptera Coleoptera Coleoptera Coleoptera Coleoptera Coleoptera Coleoptera Coleoptera Coleoptera Coleoptera Coleoptera Coleoptera Coleoptera Diptera Diptera Diptera Diptera Diptera Diptera (continued) 4. (continued) Table

BIOLOGY AND CONTROL OF EMERALD ASH BORER 36 CHAPTER 2: ECOLOGICAL IMPACTS OF EMERALD ASH BORER

Comments Comments is alternate host. host. alternate is Seed or samara gall;Seed samara or undescribed; near not species European C. marchali. Reared from seed from Reared Ontario collection from in Gagné (Raymond prep.). Abies Few Diaspididae are are Diaspididae Few specialists; confirma­ is monophagy of tion needed. Type locality about 42 locality about Type bor­ Arizona miles from if species Uncertain der. beenhas from reported Diaspididae US; few specialists (Benare pers. comm.); Normark mono­ of confirmation phagy needed. is Reference(s) Reference(s) Raymond Gagné in Gagné Raymond prep. Felt, 1918; Gagné, 1918; Gagné, Felt, 1989 1918; Gagné, Felt, 1989 1918; Gagné, Felt, 1989; Raymond in litt. Gagné 1989 Gagné, in Gagné Raymond prep. DeAngelis, 1998; DeAngelis, 2014; Carol TTD, Dohlen in litt. von TTD, 2014; Carol 2014; Carol TTD, Dohlen in litt. von Eas­ and Blackman von 1994; Carol top, Dohlen in litt. Eas­ and Blackman von 1994; Carol top, Dohlen in litt. Miller 2014a; et al., Miller Douglass in litt. Miller 2014b; et al., Miller Douglass in litt. Guild phloem phloem phloem phloem phloem phloem seed gall gall seed leaflet gallleaflet gallleaflet gallleaflet gallleaflet seed or samara gallseed samara or High High High High High High High High High High High High Risk Rating Risk Rating Confirmed Hosts ConfirmedHosts Fraxinus pennsylvanica pennsylvanica Fraxinus Fraxinus americana americana Fraxinus Fraxinus americana americana Fraxinus americana Fraxinus americana Fraxinus pennsylvanica Fraxinus Abies, Fraxinus Fraxinus Abies, F. latifolia, F. dipetala, velutina Fraxinus americana, F. F. americana, Fraxinus velutina F. nigra, Fraxinus americana americana Fraxinus Fraxinus Fraxinus velutina var. var. velutina Fraxinus t ourneyi n. sp. n. sp.

n. sp. n. sp. Scientifi c Name Name c Scientifi Contarinia Dasineura apicata ( Felt) apicata Dasineura Dasineura fraxinifolia fraxinifolia Dasineura (Felt) (Osten pellex Dasineura Sacken) tumidosae Dasineura (Felt) Dasineura Prociphilus americanus americanus Prociphilus (Walker) Prociphilus fraxinifolii fraxinifolii Prociphilus (Riley) Prociphilus pergandei pergandei Prociphilus Smith probosceus Prociphilus (Sanborn) fraxini Diaspidiotus (McKenzie) Ferris Ferris fraxini Diaspis Family Family Cecidomyiidae Cecidomyiidae Cecidomyiidae Cecidomyiidae Cecidomyiidae Cecidomyiidae Cecidomyiidae Cecidomyiidae Cecidomyiidae Aphididae Aphididae Aphididae Aphididae Aphididae Aphididae Aphididae Diaspididae Diaspididae Diaspididae Taxon Diptera Diptera Diptera Diptera Diptera Diptera Diptera Diptera Diptera Hemiptera Hemiptera Hemiptera Hemiptera Hemiptera Hemiptera Hemiptera Hemiptera Hemiptera Hemiptera (continued) 4. (continued) Table

BIOLOGY AND CONTROL OF EMERALD ASH BORER 37 CHAPTER 2: ECOLOGICAL IMPACTS OF EMERALD ASH BORER Comments Comments More study needed; oth­ study More known are sp. er Banasa beto generalists.

Reference(s) Reference(s) TTD, 2014; TTD, Schwartz Michael in litt. Donald Th omas Donald Th pers. comm. TTD, 2014; Michael 2014; Michael TTD, in litt. Schwartz 1980; TTD, Henry, 2014; Michael in litt. Schwartz 2014; Michael TTD, in litt. Schwartz 2014; Michael TTD, in litt. Schwartz 2014; Michael TTD, in litt. Schwartz 2014; Michael TTD, in litt. Schwartz 2014; Michael TTD, in litt. Schwartz 2014; Michael TTD, in litt. Schwartz 2014; Michael TTD, in litt. Schwartz 2014; Michael TTD, in litt. Schwartz 2014; Michael TTD, in litt. Schwartz 2014; Michael TTD, in litt. Schwartz 2014; Michael TTD, in litt. Schwartz 2014; Michael TTD, in litt. Schwartz Guild phloem phloem phloem phloem phloem phloem phloem phloem phloem phloem phloem phloem phloem phloem phloem phloem High High High High High High High High High High High High High High High High Risk Rating Risk Rating

Confirmed Hosts ConfirmedHosts Fraxinus americana americana Fraxinus Fraxinus velutina Fraxinus Fraxinus Fraxinus americana americana Fraxinus americana, Fraxinus pennsyl­ F. latifolia, F. vanica Fraxinus pennsyl­ F. Fraxinus, vanica americana Fraxinus americana Fraxinus Fraxinus velutina pennsylvanica Fraxinus , F. latifolia Fraxinus velutina Fraxinus F. americana, Fraxinus pennsylvanica Fraxinus pennsylvanica Fraxinus

­

(Van (Van

Th omas Th

(Knight) (Knight)

Tropidosteptes vittiscuTropidosteptes tis Scientifi c Name Name c Scientifi Banasa rolstoni & Yonke Bisulcopsallus texanus texanus Bisulcopsallus (Knight) Tropidosteptes adustus Tropidosteptes (Knight) amoenus Tropidosteptes (Reuter) brooksi Tropidosteptes Kelton canadensisTropidosteptes Duzee Van cardinalis Tropidosteptes Uhler glaber Tropidosteptes (Knight) illitus Tropidosteptes Duzee) neglectusTropidosteptes (Knight) pacificus Tropidosteptes Duzee) (Van pettiti Tropidosteptes Reuter populi Tropidosteptes (Knight) venosus rufi Tropidosteptes (Knight) tricolor Tropidosteptes Duzee Van Miridae Family Family Pentatomidae Miridae Miridae Miridae Miridae Miridae Miridae Miridae Miridae Miridae Miridae Miridae Miridae Miridae Miridae Taxon Hemiptera Hemiptera Hemiptera Hemiptera Hemiptera Hemiptera Hemiptera Hemiptera Hemiptera Hemiptera Hemiptera Hemiptera Hemiptera Hemiptera Hemiptera Hemiptera Hemiptera Hemiptera (continued) 4. (continued) Table

BIOLOGY AND CONTROL OF EMERALD ASH BORER 38 CHAPTER 2: ECOLOGICAL IMPACTS OF EMERALD ASH BORER

and records

Corylus,

records likely likely records Comments Comments Hamamelis

Carya ovata, Carya ovata, and in this species likely for Henry in (Tom error litt.). Ceanothus, Populus non-Oleaceae other (Tom in error hosts Henry in litt.). Forestiera species another to refer Henry in litt.). (Tom Pecan record is doubtful doubtful is record Pecan in litt.). Smith (David Validity of this taxon this taxon of Validity (molecular) in need of verification. Only known from a from known Only in Central counties few evidently California and declining.

Reference(s) Reference(s) Usinger, 1945; Usinger, , Ruhoff and Drake Henry 1965; Tom Chionanthusin litt; Laura from record Miller in litt. Usinger, 1945; Usinger, , Ruhoff and Drake Henry 1965; Tom in litt. Froeschner, 1944; Froeschner, 1945; Usinger, , Ruhoff and Drake Henry 1965; Tom Miller Laura in litt.; in litt Williams, 2007; Williams, in litt. Smith David David Smith in litt. in litt. Smith David Langford and 1935; McConnell, in litt. Smith David 2005, 2007; Wagner, et al., Robinson 2014 2002 et al., Wagner DLW unpubl. data data unpubl. DLW 1970; Wag­ Buckett, 2007 ner, leaves leaves leaves leaves Guild phloem phloem phloem new leaves new leaves new leaves new leaves new leaves new leaves mature leaves leaves mature leaves mature High High High High High High Very High High Very Risk Rating Risk Rating High to Moderate Moderate to High High to Moderate Moderate to High High to Moderate Moderate to High High to Moderate Moderate to High

and other other and

Confirmed Hosts ConfirmedHosts Chionanthus, Fraxinus Fraxinus Chionanthus, Fraxinus latifolia, F. F. latifolia, Fraxinus velutina, Oleaceae Chionanthus, Fraxinus Fraxinus Chionanthus, americana Fraxinus americana, americana, Fraxinus F. latifolia, F. nigra, F. pennsylvanica Fraxinus americana americana Fraxinus Fraxinus americana, F. F. americana, Fraxinus latifolia , Syringa Fraxinus americana Fraxinus Fraxinus gooddingii Fraxinus latifolia latifolia Fraxinus

(Hy. (Hy.

(Say) (Say)

) McAtee McAtee

(Say) (Say)

inornata near

Scientifi c Name Name c Scientifi Leptoypha costata Parshley Leptoypha minor nubilis(including Leptoypha mutica Eupareophora parca (Cresson) Tethida barda barda Tethida Tomostethus multicinctus Tomostethus (Rohwer) Olceclostera angelica Olceclostera (Grote) Hydrelia Philtraea elegantaria Edwards) Philtraea latifoliae Buckett Tenthredinidae Tenthredinidae Family Family Tingidae Tingidae Tingidae Tingidae Tingidae Tingidae Tenthredinidae Tenthredinidae Tenthredinidae Tenthredinidae Apatelodidae Apatelodidae Geometridae Geometridae Geometridae ­ ­ ­ Taxon Hemiptera Hemiptera Hemiptera Hemiptera Hemiptera Hemiptera Hymenop tera Hymenop tera Hymenop tera Lepidoptera Lepidoptera Lepidoptera Lepidoptera Lepidoptera Lepidoptera Lepidoptera (continued) 4. (continued) Table

BIOLOGY AND CONTROL OF EMERALD ASH BORER 39 CHAPTER 2: ECOLOGICAL IMPACTS OF EMERALD ASH BORER

(Charley (Charley as a host a host as record from in Big Bend in Big NP Comments Comments F. greggii F. TX Mc­ (Noel Laredo, pers. comm.) Farland DLW; Bend Big by and Lycium unpubl. Wagner from two possibly but data in which taxa involved case feeder is the ash imperiled . is listed in error in in error listed is tabulations; previous C. traces to record that cucullipennella in litt.). Eiseman Ligustrum Found along creeks in creeks along Found northern and central California. Currently known only only known Currently York. New Upstate from cell layer single on Feeds cambium. in cork Currently known only only known Currently York. New Upstate from cell layer single on Feeds initial­ cambium in cork into then tunnels and ly layers. deeper cork Feeds on single cell layer cell layer single on Feeds cambium. in cork

Reference(s) Reference(s) Noel McFarland Noel Wag­ pers. comm.; data unpubl. ner Buckett, 1970; Wag­ Buckett, 2007 ner, 2007 Wagner, Johnson and Lyon, Lyon, and Johnson et al., 1988; Pohl 2007 2004; Wagner, Wagner, 2007, Wagner, unpubl. data Fitzgerald, 1973; Fitzgerald, 2007 Wagner, Fitzgerald, 1973; Fitzgerald, 2007 Wagner, Fitzgerald and and Fitzgerald 1971; Simeone, 1973; Fitzgerald, 2007 Wagner, Schmidt et al., 2014 et al., Schmidt leaves leaves leaves leaves leaves leaves Guild crown new leaves new leaves new leaves new leaves mature leaves leaves mature bark miner that that miner bark tunnels into root root into tunnels green bark miners miners bark green green bark miners miners bark green High High High High High High Very High High Very Very High High Very comments comments Risk Rating Risk Rating Moderate but see but Moderate

latifolia latifolia

Confirmed Hosts ConfirmedHosts and greggii Fraxinus Lycium Fraxinus dipetala dipetala Fraxinus Fraxinus americana, F. F. americana, Fraxinus quadrangulata F. americana, Fraxinus pennsylvanica F. nigra, Fraxinus Fraxinus Fraxinus pennsylvanica pennsylvanica Fraxinus Fraxinus pennsylvanica pennsylvanica Fraxinus Fraxinus americana, F. F. americana, Fraxinus pennsylvanica Fraxinus velutina

Fitz­

n. sp. n. sp.

Scientifi c Name Name c Scientifi Philtraea paucimacula paucimacula Philtraea McDunno­ Barnes and ugh Philtraea surcaliforniae Buckett Plagodis kuetzingi Plagodis (Grote) (Ely) fraxinella Caloptilia Caloptilia Marmara basidendroca Marmara Fitzgerald Marmara corticola Marmara gerald Marmara fraxinicola Marmara Braun Chloronycta tybo tybo Chloronycta (Barnes) Family Family Geometridae Geometridae Geometridae Gracillariidae Gracillariidae Gracillariidae Gracillariidae Gracillariidae Noctuidae Noctuidae Taxon Lepidoptera Lepidoptera Lepidoptera Lepidoptera Lepidoptera Lepidoptera Lepidoptera Lepidoptera Lepidoptera Lepidoptera Lepidoptera Lepidoptera Lepidoptera Lepidoptera Lepidoptera Lepidoptera Lepidoptera (continued) 4. (continued) Table

BIOLOGY AND CONTROL OF EMERALD ASH BORER 40 CHAPTER 2: ECOLOGICAL IMPACTS OF EMERALD ASH BORER

­

as a as

records likely likely records slight slight syringae: ; reports from from ; reports

Comments Comments (Zacharczenko et (Zacharczenko hosts are are hosts Fraxinus should be verified. should

Non- inciden­ insignificant, (Eric erroneous tal, or Larvae in litt.). Quinter in vigorous feed only usually shoots growing saplings. of Triosteum S. to refer and in error forbesi 2014). al. CDFA record for Keck for record CDFA antirrhinoides iella from derives likely host of S. a misidentification picina F. greggii F. of leaves New in Chisos Ba­ growing Spring Bendsin, Big NP. active. Sister to gen- period flight and COI erences; diff italic with shared barcode syringae Olea DLW discounts all discounts DLW those than other records Oleaceae: Fraxinus on host. preferred

Reference(s) Reference(s) Wagner, 2007; Rob­ Wagner, 2014 et al., inson 2007; Rob­ Wagner, 2014 et al., inson Wagner, 2007; Wagner, 2011; et al., Wagner et al., Robinson 2014 Robinson et al., et al., Robinson card 2014; CDFA le database pest fi Crumb, 1956; Crumb, le pest fi card CDFA database DLW unpubl. data data unpubl. DLW Robinson et al., et al., Robinson 2014 Eichlin and Duck- and Eichlin 1988; Wag­ worth, 2007; Robinson ner, 2014 et al., Tuttle, 2007; Tuttle, 2005, 2007 Wagner, 2007; Tuttle, 2005, 2007 Wagner, leaves leaves Guild xylem (borer) (borer) new leaves new leaves new leaves new leaves new leaves new leaves new leaves new leaves new leaves new leaves mature leaves leaves mature mature leaves leaves mature new shoot feeder new shoot High High High High High High High Risk Rating Risk Rating High to Moderate Moderate to High High to Moderate Moderate to High High to Moderate Moderate to High , ,

, Syringa , F. nigra , and Confirmed Hosts ConfirmedHosts Fraxinus americana americana Fraxinus Fraxinus americana, F. F. americana, Fraxinus pennsylvanica F. nigra, Chionanthus virginicus, virginicus, Chionanthus F. americana, Fraxinus pennsylvanica F. nigra, Fraxinus, F. latifolia latifolia F. Fraxinus, Fraxinus, F. velutina velutina F. Fraxinus, Fraxinus cuspidata cuspidata Fraxinus Fraxinus americana, F. F. americana, Fraxinus pennsylvanica F. nigra, Fraxinus, F. americana, americana, F. Fraxinus, Olea pennsylvanica, F. Fraxinus velutina pennsylvanica F. Ligustrum Chionanthus virginicus , american Fraxinus caroliniana F. ­

(J. E. (J.

(J. B. B. (J. (Grote) (Grote) Scientifi c Name Name c Scientifi Copivaleria grotei Mor grotei Copivaleria rison furcata Smith) Sympistis chionanthi Smith) Smith) Sympistis fortis Sympistis punctilinea (Hampson) Sympistis heterogena Sympistis (A. Blanchard) Palpita magniferalisPalpita (Walker) Podosesia aureocincta Podosesia & Nielson Purrington Ceratomia sonorensis Hodges Ceratomia undulosa (Walker) Family Family Noctuidae Noctuidae Noctuidae Noctuidae Noctuidae Noctuidae Noctuidae Noctuidae Noctuidae Noctuidae Noctuidae Noctuidae Pyralidae Sesiidae Sphingidae Sphingidae Sphingidae Sphingidae Taxon Lepidoptera Lepidoptera Lepidoptera Lepidoptera Lepidoptera Lepidoptera Lepidoptera Lepidoptera Lepidoptera Lepidoptera Lepidoptera Lepidoptera Lepidoptera Lepidoptera Lepidoptera Lepidoptera Lepidoptera Lepidoptera Lepidoptera Lepidoptera (continued) 4. (continued) Table

BIOLOGY AND CONTROL OF EMERALD ASH BORER 41 CHAPTER 2: ECOLOGICAL IMPACTS OF EMERALD ASH BORER

likely the likely traces to pre­ traces to

Comments Comments (Chris S. mexicana

DLW discounts all discounts DLW those than other records Oleaceae. on DLW discounts all discounts DLW those than other records Oleaceae. another on species likely Fraxinus host beto principal Ohio of species south River. DLW discounts all discounts DLW those than other records Oleaceae. on DNA barcodes sug­ barcodes DNA Arizona that gestive S. chersis of populations sister n. sp. represent to in litt.) Schmidt DLW discounts all discounts DLW those than other records Oleaceae; already on declin­ and uncommon preferred Fraxinus ing; host. DLW discounts all discounts DLW those than other records Oleaceae. on Forestiera host. principal Report by Wagner Wagner Report by Z.(2007) as hepariella ; the use of in error hepariella liminary determination (in litt.); Eichlen Tom by still under taxonomy study.

Reference(s) Reference(s) Tuttle, 2007; Tuttle, 2005, 2007 Wagner, Tuttle, 2007; Tuttle, 2005, 2007 Wagner, Tuttle, 2007; Tuttle, 2005, 2007 Wagner, Chris in Schmidt litt. Tuttle, 2007; Tuttle, 2005, 2007 Wagner, Tuttle, 2007; Tuttle, 2005, 2007 Wagner, Tuttle, 2007 Tuttle, Wagner, 2007 Wagner, leaves leaves Guild mature leaves leaves mature mature leaves leaves mature mature leaves leaves mature mature leaves leaves mature mature leaves leaves mature mature leaves leaves mature mature leaves leaves mature High High High Moderate Moderate Very High High Very Very High High Very Risk Rating Risk Rating High to Moderate Moderate to High High to Moderate Moderate to High

­

spp., Frax spp.,

latifolia latifolia

(preferred host) host) (preferred

Confirmed Hosts ConfirmedHosts , Syringa Fraxinus Fraxinus americana americana Fraxinus this host of (origin F. verification) requires nigra Fraxinus americana, americana, Fraxinus pennsylvanica, F. Ligustrum, velutina, F. Oleaceae other Syringa, Fraxinus velutina Fraxinus americana, americana, Fraxinus Ligustrum Chionanthus virginicus, virginicus, Chionanthus F. ameicana, Fraxinus pennsylvanica, F. nigra, Ligustrum, Syringa Forestiera inus gooddingii, F. greggii Fraxinus Fraxinus

hepariella hepariella chersis

near

Scientifi c Name Name c Scientifi Manduca jasminearum (Guérin) Sphinx canadensis canadensis Sphinx Boisduval (Hübner) (Hübner) chersis Sphinx Sphinx near Sphinx Sphinx franckii franckii Sphinx Neumoegen J. E. J. kalmiae Sphinx Sphinx libocedrus Zelleria Family Family Sphingidae Sphingidae Sphingidae Sphingidae Sphingidae Sphingidae Sphingidae Sphingidae Sphingidae Sphingidae Sphingidae Sphingidae Sphingidae Sphingidae Yponomeutidae Taxon Lepidoptera Lepidoptera Lepidoptera Lepidoptera Lepidoptera Lepidoptera Lepidoptera Lepidoptera Lepidoptera Lepidoptera Lepidoptera Lepidoptera Lepidoptera Lepidoptera Lepidoptera Lepidoptera (continued) 4. (continued) Table

BIOLOGY AND CONTROL OF EMERALD ASH BORER 42 CHAPTER 2: ECOLOGICAL IMPACTS OF EMERALD ASH BORER Comments Comments A similar appear­ A similar occurs moth in ing California on southern Forestiera. Reference(s) Reference(s) John DeBenedictisJohn unpubl. data leaves leaves Guild High? High? Risk Rating Risk Rating Confirmed Hosts ConfirmedHosts Fraxinus dipetala dipetala Fraxinus semitincta

near

Scientifi c Name Name c Scientifi Zelleria Family Family Yponomeutidae Taxon Lepidoptera Lepidoptera (continued) 4. (continued) Table

BIOLOGY AND CONTROL OF EMERALD ASH BORER 43 CHAPTER 2: ECOLOGICAL IMPACTS OF EMERALD ASH BORER

Figure 22. Mating pair of North America’s largest beetle, Dynastes granti (Photo courtesy Margarethe Brummerman). So far as known, actively growing ash shoots are the principal feeding site for this behemoth, which sometimes exceeds 80 mm in length. Males chew a lesion into new growth and females are paired as they arrive at the wound. Curiously, its more widespread cousin, the Eastern ; Dynastes tityus, is virtually unstudied as an adult. The only adult feeding record of which we are aware, again is for a Fraxinus (Glaser 1976). Grant’s rhinoceros beetle is quite sought-after by photographers and collectors; persons (even families) with interests in Coleoptera, sometimes travel from as far away as Japan and Korea to see the beetle. See also caption for Figure 23.

species as Very High (n = 2), High (n = 3), High attention-worthy insects may be threatened by EAB: to Moderate (n = 3), and Moderate (n = 1). Three the western rhinoceros beetle (Dynastes granti), of the nine have restricted global distributions and whose adults occasionally exceed 80 mm in length are ranked as G4 species in NatureServe Explorer (Figs. 18, 22). While this large scarab is a generalist (2014): Manduca jasminearum, Sphinx canadensis, as a larva, feeding in dead wood of various tree and S. franckii (Fig. 16). The first of these is already species, its adult may prove to be a specialist that in decline along the northern portion of its range feeds primarily on the new growth of velvet ash (NatureServe Explorer, 2014), and sphingids (and in Arizona. Courting males chew into the cambial especially members of the nominate genus Sphinx) layers of an ash tree and then await the arrival of have been identified as a group in decline across females (which presumably feed and mate at the much of the northeastern United States, perhaps wound site). The best known and most reliable due to the exotic tachinid Compsilura concinnata locations for the beetle occur in the vicinity of velvet (Meigen) (Wagner, 2012) or an as yet unidentified ash, where the adults have access to vigorously parasitoid (Sam Jaffe, unpublished data). growing green shoots (Margarethe Brummerman, One of the most surprising findings from our Patrick Sullivan, and Bill Warner, all in litt.). Beetle study is that one of North America’s largest, and collectors visit the southwestern United States with

BIOLOGY AND CONTROL OF EMERALD ASH BORER 44 CHAPTER 2: ECOLOGICAL IMPACTS OF EMERALD ASH BORER the Grant’s rhinoceros beetle as a common focus of their itineraries. Rhinoceros beetles are also popular pets in some Asian countries; we know of instances of Asian families traveling to Arizona primarily to acquire living Dynastes adults to keep as pets and captive breeding (Fig. 23). Dynastes granti is even occasionally sold in Japanese pet shops and has appeared in staged beetle fights. Less is known about the habits and dietary proclivities of the eastern rhinoceros beetle (Dynastes tityus), but it too is known to chew into shoots of ash and feed at the wounds (Glaser, 1976); no other adult food plants are known to us. Ratcliffe (2009) called Figure 23. Entomophily. The insect zoo at the United States attention to the threat of EAB to two other rhinoceros Museum of Natural History, ranks only behind dinosaurs beetles in the genus Xyloryctes, which reach lengths as a public draw. Insects also enjoy considerable favorable close to 30 mm (Fig. 20). Both North American attention in many Asian countries. This photo is from an insect exhibition held in the Tokyo Tower over a six-week span in 2013 species are specialists on ash roots as larvae. If even a (http:www.japantrends.com/tokyo-tower-insect-exhibition/) few of the species mentioned above were to disappear, (Photo courtesy of Tokyo Tower, Nippon Television City Corporation). More than 16,000 insects were on display, many the losses to North American arthropod fauna would of which were alive in the featured insect jungle. The scene be great (in both senses of the word). above shows a beetle enclosure with many horned scarabs, kindred to and perhaps even including the North American Dynastes and Xyloryctes that are listed in Table 4 and shown in Figures 18, 20, and 22. Asian families are also known to travel annually to the Southwest United States to look for live D. granti adults to be kept as family pets and bred.

Figure 24. Feeding guilds of specialist herbivores on ash. Chart shows 98 species of ash specialists divided by feeding guild: bark (3), phloem (26), cambium (9), xylem (2), root (4), seed (8), new shoot feeder (3), gall-former (10), leaf miner (2), leaves (32), and unknown (1). Some species could be included into two feeding guilds, e.g., gall formers may form galls on seeds or leaves, etc. In these cases, we grouped individuals into the most applicable guild. See Table 4 for additional notes on feeding guilds.

BIOLOGY AND CONTROL OF EMERALD ASH BORER 45 CHAPTER 2: ECOLOGICAL IMPACTS OF EMERALD ASH BORER

Feeding Guilds and Specificity of Arthropod Herbivores

We group the 98 phytophagous arthropods identified in Table 4 into ten feeding guilds (Fig. 24). The degree of host specificity for the majority of the species listed in Table 1 is not known in detail. We are unaware of any taxonomically comprehensive study of ash herbivores where researchers thoroughly sampled across the spectrum of available Fraxinus (and other Oleaceae). It is our belief that most Fraxinus specialists (as defined in this work) consume a spectrum of available ash species Figure 25. Marmara fraxinella, Family Gracillariidae (Photo courtesy Charley Eiseman). Leafminers of the genus Marmara growing in a given locale. Stated differently, we see little are specialized herbivores. Three North American Marmara indication that the phylogenetic, physiological, and are known only to feed on Fraxinus, and two of these mine morphological differences across the genus Fraxinus only in the bark of young, actively growing ash trees. are so great that ash-specialized herbivores are routinely restricted to a single species or even Section of the genus (contrary to what one might be led to believe from published host records). In the western United States many of the herbivores listed in Table 4 are associated with a single ash species simply because only one Fraxinus (or member of the Oleaceae) occurs at a given locale. In eastern North America, in cases where only Fraxinus (without a species indicated) is listed in Table 4, the host is likely assignable to F. americana and/or F. pennsylvanica. Based on our literature review, communications with >80 taxonomists, and DLW’s 30­ Figure 26. Four males of the two-tailed swallowtail (Papilio year rearing program of North American Lepidoptera multicaudata), Arizona’s state butterfly, puddling in Oak Creek canyon, outside of Sedona, Arizona. Larvae feed on members (that has encompassed more than 2,000 species of three plant families, but only ashes are present in some of representing more than 50 families, including miners, the western canyons where this large swallowtail butterfly occurs. (Photo courtesy Tyger Gilbert, www.TygerGilbert.com) gall-formers, wood feeders, and others), the following generalities emerged. 4 will be found to feed on multiple species of ash. The Wood feeders. As reflected in our results, wood aphids (n = 3), diaspidid scales (n = 2), and mirids feeders (restricted here to taxa consuming mostly (n = 15) appear to be obligate Fraxinus feeders. Tom subcambial tissues including dead and live wood Henry (in litt.) believes that the three tingids listed feeders) are seldom specialized in diet, e.g., not one in Table 4 are oligophagous on Oleaceae and that of North America’s 1000 cerambycids is known to be contrary host records are unreliable. The diet breadth a Fraxinus specialist as a larva. We list only two wood of the pentatomid B. rolstoni requires further study feeders: the buprestid, Trigonogya reticulaticollis, as very few stink bugs are known to be host plant whose host range is not well investigated and the specialists. clearwing borer Podosesia aureocincta, which is likely Cambium feeders. We record seven bark beetles oligophagous on Oleaceae. and a cambium-mining agromyzid (Phytobia sp.) as Phloem feeders. We identify 25 Fraxinus ash specialists. The only buprestid identified as an specialist, phloem-feeding hemipterans. While several ash specialist – from a family with more than 760 of these appear to be monophagous, we believe that North American species – was Agrilus subcinctus, a this is a sampling artifact and that the species in Table cambium miner (of course, we exclude A. planipennis).

BIOLOGY AND CONTROL OF EMERALD ASH BORER 46 CHAPTER 2: ECOLOGICAL IMPACTS OF EMERALD ASH BORER

As a subcategory to cambium feeders, we include the its dependency on Oregon ash is likely a geographical three gracillariids that mine in green bark. Marmara artifact because its host is the only member of the basidendroca tunnels into cambial layers at least for Oleaceae that grows where the moth occurs. One part of its life cycle (Fitzgerald, 1973). This gracillariid surprising outcome of our compilation is how and two of its congeners, M. corticola and M. proportionately few specialist external leaf feeders fraxinicola are all highly specialized insects regarded to were detected (<32% of the 98 Fraxinus-dependent be at high risk (Fig. 25). herbivores), though this guild comprises most of the New shoot feeders. We include the borer insect biomass using ash and the most conspicuous Papaipema furcata (Noctuidae), which so far as is herbivores on Fraxinus. Clearly, the insects that live known, tunnels only in new growth of Melioides and feed inside seeds, stems, or leaves dominate our Fraxinus. Both western and eastern rhinoceros list of species threatened by EAB. beetles (D. granti and D. tityus) can be provisionally placed here because adults are only known to feed at Oligophages, Polyphages, and Other Trophic Levels vigorously growing shoots of ash. Root borers. We report four coleopterans whose Our ecological assessment focused on specialists; it larvae are ash root specialists – two leaf beetles: ignored oligophages and polyphages that might be Trichaltica tibialis and T. scabricula and two scarab affected by ash decline. However, over the course beetles: Xyloryctes jamaicensis and X. thestalus (Fig. of preparing this manuscript, we discovered a few 20). The extent to which the larvae of these beetles instances where an oligophage not treated in our list accept different species of ash has not been studied, might be affected by EAB. For example, the two-tailed although their ranges are broad enough to suggest that swallowtail (Papilio multicaudatus) (Fig. 26) uses none is monophagous. Fraxinus, Prunus, and Ptelea as hosts in arid lands

Seed feeders. Members of this guild, or at least of western North America, but only ash species are the weevils of the genus Lignyodes (Fig. 19), appear to available in some canyons where this butterfly lives. be oligophagous. Psomus armatus, so far known only Octotoma (hispine chrysomelids), e.g., O. plicatula from white ash, is expected to feed on other ashes and O. marginicollis, were not included because their based on what is known of the diet breadth of related leafmining larval stages are specialists on non-ash weevils. species, even though the adult beetles are believed to Gall formers. Gall insects are widely recognized to do much of their adult feeding on Fraxinus species be among the most specialized insect herbivores (Felt, (Shawn Clark, in litt.). 1918, Gagné, 1989). The two gall-forming eriophyid We did not find mention of any specialist mites that we list have been reported from more than parasitoids, predators, or pathogens that we could one species of Section Melioides ashes. Gagné (in litt.) confidently state were dependent on an ash herbivore. believes that all the cecidomyiid gall formers in Table 1 The recently described Mymaromella pala Huber will be found to occur on more than one species within & Gibson (Mymarommatidae) is known only from a Fraxinus Section, but that significant differences are ash log collections, but its presumed host is a bark- likely to exist among Sections. residing psocid (Huber et al., 2008) likely to dwell on Leafminers. There are only two leafminers in other trees as well. Gagné (1989) regards Contarinia our study, both members of the genus Caloptilia thalactri to be a phytophagous inquiline in the galls of (Gracillariidae). One leafminer, Caloptilia n. sp., is Dasineura tumidosae. However, beyond this inquiline an ecological specialist on F. latifolia; its eastern sister record, we did not find mention of any other taxon, C. fraxinella, feeds on at least three species indirectly ash-specialized insects. from two Fraxinus sections. External (chewing) leaf feeders. Only one externally feeding lepidopteran is recorded as monophagous: Philtraea latifoliae (Geometridae), and

BIOLOGY AND CONTROL OF EMERALD ASH BORER 47 CHAPTER 2: ECOLOGICAL IMPACTS OF EMERALD ASH BORER

disCUssion miners known only from a restricted area in Upstate New York. Philtraea latifoliae (Geometridae) feeds Imperilment Risk Ratings exclusively on Fraxinus latifolia and is known only from a few counties in central California (Buckett, It is our estimate that no less than 98 species of 1970). Sphinx canadensis’ rating was raised to Very invertebrate herbivores would be appreciably affected High because it is principally associated with F. nigra, by an elimination or massive reduction in abundance an ash that is highly susceptible to EAB infestation of North America’s 16 native Fraxinus. For these (Leah Bauer pers. comm.) and which is predicted to herbivores we estimate endangerment risks as Very be increasingly at risk due to climate change (Liang High (n = 5), High (n = 75), High to Moderate (n = and Fei, 2014). Sphinx franckii is already uncommon 15), and Moderate (n = 3) (Fig. 21). Forty-five of the and northern populations are in decline (Wagner, species-level taxa listed in Table 4 are identified as 2012). “at risk” for the first time. Eighteen (6%) of the taxa For perspective on the importance of ash and identified by Gandhi and Herms (2010) had their the emerald ash borer, it is useful to compare ash imperilment status upgraded; twenty six (35%) of decline with what is known about American chestnut the species listed in their treatment as being of high (Castanea dentata [Marsh.] Borkh.) and its dependent to moderate endangerment risk are dropped from herbivore fauna. Opler (1978) listed seven species of consideration (Appendix 3). A key message deriving Lepidoptera that may have gone extinct as a result from this expert-based approach is that researchers of losing this once ecologically dominant forest tree. should use caution when gleaning host records from One of these seven moths has since been rediscovered dated literature, the internet, and especially secondary (Synanthedon castaneae [Busck]) (Anagnostakis et al., and tertiary resources. We found records (and 1994), and another listed species, Tischeria perplexa especially older host records) to be rife with invalid Braun, may not be a valid species given that other taxa, erroneous identifications, instances where a Fagaceae-feeding Tischeria from the eastern United presumed ash specialist was not phytophagous, cases States are not chestnut specialists (Braun, 1972)4. where presumed ash specialists were generalists (type Thus, five is a better estimate of the number of moth I errors), cases where presumed oligophagous or species that have been lost from the North American generalist herbivores were in fact specialists (type II fauna5 due to American chestnut decline. If one errors), and cases where exotic or extralimital taxa compares the number of extinct moths restricted to were included. chestnut (n = 5) to the at risk ash-feeding Lepidoptera As noted previously, we discounted the ecological listed in Table 4 (n = 32), one is immediately struck by importance of introduced (exotic) plants, e.g., lilac, the magnitude of the threat posed by the introduction privet, and olive, as alternative hosts that could and spread of EAB. North America has not faced a support sufficiently large populations to ensure a threat of this magnitude to its native insect herbivore taxon’s long-term survival in North America. This biodiversity from an exotic species over the course of seems to be a defensible position if one were to invoke the last two centurie. We believe that this is because the precautionary principle in assessing risk. But, we also recognize that it is possible that some invasive 4 Three other Tischeria occur on Castanea dentata – one of plants (such as Ligustrum) could play a role in the which, T. castaneaeella, is believed to be quite closely related to T. perplexa, the purported Castanea dentata specialist. All three survival of some native Oleaceae specialists were EAB of the extant Tischeria that fed on chestnut are breeding on to eliminate much of the Fraxinus in a region. red oaks. Given the above, it is DLW’s belief that T. perplexa will The Very High risk rating – given to just five prove to be a taxonomic synonym. 5 As a caveat, it is worth noting that no one has systematically lepidopterans – was reserved for ash specialists sampled introduced Castanea or related native Castanea, and believed to be at risk or in decline due to other especially Alleghany chinquapin (C. pumila), which co-occurred with American chestnut; nor have workers surveyed nut-produc- causes. Marmara basidendroca and M. corticola ing stands of C. dentata in Ohio, Maine, and elsewhere for the (both Gracillariidae) are both specialized stem missing species.

BIOLOGY AND CONTROL OF EMERALD ASH BORER 48 CHAPTER 2: ECOLOGICAL IMPACTS OF EMERALD ASH BORER

Fraxinus represents a phylogenetically and or revision, i.e., some names in Table 4 may prove to be chemically isolated biological island that is too far synonyms (Michael Schwartz pers. comm.). removed from its allied genera (e.g., Chionanthus, In summary, we recognized 98 species of Forestiera, exotic Ligustrum, and Oleaeceae) for herbivores as being threatened by the loss of these plants to serve as refuges for Fraxinus specialist Fraxinus in the United States and Canada. No herbivores over ecological and evolutionary time. doubt, additional ash specialists will be identified in the coming decades. Knowledge of herbivores Data Gaps feeding on ash is severely lacking in the southern and western United States; very little is known of Host ranges for virtually all of the taxa in this study the phytophagous insects feeding on six western remain incompletely known. Targeted herbivore Fraxinus species (F. anomala, F. cuspidata, F. dipetala, surveys of the 16 North American Fraxinus would be F. gooddingii, F. greggii, and F. papillosa). Gall midges valuable, especially for understudied ash species and (Cecidomyiidae), curculionid weevils, mites, thrips, those currently under threat from EAB or soon to be and other taxonomically challenging arthropods can attacked as the beetle’s range expands. In particular, be expected to yield additional Fraxinus specialists. southern ash are poorly studied as they tend to grow The latter two taxa seem especially likely to include in swamps and all nine western ash species require ash specialists because their taxonomy is nascent further research; almost nothing is known of F. and no systematic continent-wide surveys have been albicans, F. berlandieriana, F. cuspidata, F. gooddinggii, carried out. Across all taxa, molecular markers can be and F. papillosa. Likewise, the importance of other expected to reveal new cryptic specialist herbivores, native Oleaceae, especially Forestiera, as hosts for ash- especially in those taxa where the species-level feeding herbivores is in need of study. Of critical taxonomy has proven difficult. importance to the evaluation of imperilment risk is the possibility that other native Oleaceae may be susceptible to the emerald ash borer. Only very ConClUsion recently did researchers learn that Chionanthus is also susceptible to attack (Entomology Today, 2014). For the purposes of our ecological assessment we Other Oleaceae species such as Osmanthus Lour., a embraced the precautionary principle. The body of genus of trees found in black water streams of the our assessment is, at its essence, a doomsday scenario: southern United States, remain virtually unstudied. what might transpire if North America lost all of While the susceptibility of other native Oleaceae to its Fraxinus or if ash numbers dwindled to a point EAB is of no commercial relevance, it is a matter of where ash lost functional value in North American considerable conservation significance. wildlands. Presently, EAB is spreading at a dramatic We suspect that modern systematic analyses pace. We hope a growing number of parasitoids, employing molecular markers will reveal additional pathogens, and predators will soon reduce the beetle’s cryptic species that are ash specialists (especially hyperabundance and rate of spread, and that some in the western United States where species-level subset of ash species, genotypes, age classes, etc. will taxonomic studies lag). One interesting example prove resistant to EAB. However, should the beetle that surfaced over the course of our studies involves cause catastrophic losses of ash, as many as 150 U.S. the sizable and charismatic great sphinx (S. chersis). plant community types (16 of which are regarded as COI barcodes suggest the hawkmoth is two species, imperiled or critically imperiled) could be severely with the populations from southeastern Arizona compromised. Fraxinus has a surprisingly rich, representing an as yet undescribed species that specialized, beautiful, and noteworthy invertebrate Chris Schmidt (in litt.) believes is closely related to fauna, including hercules beetles, rhinoceros S. mexicana. Conversely, some taxa in our treatment beetles, and hawkmoths – some of our continent’s may have been over split taxonomically. The mirid most magnificent invertebrates. By our assessment, genus with 14 Tropidosteptes species is in need of the magnitude of North American invertebrate

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biodiversity loss could greatly exceed that associated Hayes, Bill Johnson, Ken Metzler, Mike Quinn, with American chestnut blight. The number of Jim McCormac (3), Tom Murray, Anton Reznicek plant communities likely to be affected, the number (2), Judy Semroc (4), and Tokyo Tower (Nippon of herbivores at risk (nearly 100 species), and the Television City Corporation). Virge Kask helped with charismatic nature of the fauna in peril, argue for the preparation of figures. Support for this work, in continued and increased efforts to bring the emerald part, was provided by USDA Forest Service FHTET ash borer under control. Co-op Agreement 14-CA-11420004-138. Additional assistance came from a State Wildlife Grant from the aCKnoWledgments Wildlife Division of the Connecticut Department of Environmental Protection. This work is a compilation of the collective knowledge of more than 100 biologists, collectively reFerenCes representing >2500 person-years of accumulated study and knowledge, who made their expertise, Anagnostakis, S. L., K. M. Welch, J. W. Snow, K. and in some cases unpublished data, available to Scarborough, and T. D. Eichlin. 1994. The us. It was an honor to work with this august group. rediscovery of the clearwing chestnut moth, Botanists, forest managers, ecologists, and wildlife Synanthedon castaneae (Busck) (Lepidoptera: biologists – all of which are listed by name – are Sesiidae) in Connecticut. Journal of the New York given in Appendices 1 and 2. Anton Reznicek, Jim Entomological Society 102: 111–112. Bissell, Jim McCormac, and Judy Semroc greatly Anulewicz, A. C., D. G. McCullough, and D. L. shaped the content of what we present with regard to Cappaert. 2007. Emerald ash borer (Agrilus effects on plant communities and the likely cascading planipennis) density and canopy dieback in three consequences of emerald ash borer. Principal North American ash species. Arboriculture and authorities and sources for specialist herbivores are Urban Forestry 33: 338–349. listed in Table 4. We are especially indebted to those Anulewicz, A. C., D. G. McCullough, D. L. Cappaert, who waded through multiple emails and reviewed and T. M. Poland. 2008. Host range of the preliminary lists. Roughly in order of deceasing emerald ash borer (Agrilus planipennis Fairmaire) email correspondence: Ray Gagné (Cecidomyiidae), (Coleoptera: Buprestidae) in North America: John DeBenedictis (Yponomeutidae), Margarethe results of multiple-choice field experiments. Brummerman (Dynastes), Tom Henry (Hemiptera), Environmental Entomology 37: 230–241. Rob Huber (Sesiidae), Charley Eisemen (leafminers, Arnett, R. H. Jr. 2000. American Insects: A Handbook weevils), Jim Tuttle (Sphingidae), Charlie and of the Insects of America North of Mexico. Second Lois O’Brien (Curculionidae), Robert Anderson Edition. CRC Press, Boca Raton, Florida, USA. (Curculionidae), Michael Schwartz (Miridae), 1024 pp. Rick Westcott (Coleoptera), Terry Fitzgerald Arnett, R. H., Jr., M. C. Thomas, P. E. Skelley and J. H. (Gracillariidae), Shawn Clark (Chrysomelidae), Frank (eds.). 2002. American Beetles, Volume II: David Smith (Tenthredinidae), and Laura Miller : Scarabaeoidea through Curculionoidea. (Tingidae). Several colleagues and collectors CRC Press LLC, Boca Raton, Florida, USA. 880 pp. supplied unpublished observations or data, especially Atkinson, T. H. 2014. Bark and ambrosia beetles of Margarethe Brummerman, Ted MacRae, and Bill Canada and the U.S. (excluding Hawaii). Complete Warner. Roy Van Driesche, Faith Campbell, Tate URL (http://www.barkbeetles.info/us_canada_ Lavitt, Virginia Wagner, and David Cappaert made chklist_expanded.php). numerous helpful suggestions on drafts of the Baker, E. W. and Y. Suigong, 1988. A catalog of the manuscript. Images were supplied by Daniel Boone, false mites (Tenuipalpidae: Acari) of the Margarethe Brummerman (2), Charley Eiseman, United States. International Journal of Acarology Tyger Gilbert (www.tygergilbert.com), Richard 14: 143–155.

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Morgantown, West Virginia, USA. New Jersey, USA. 512 pp. Slesak, R. A., C. F. Lenhart, K. N. Brooks, A. W. Wagner, D. L. 2007. Emerald ash borer threatens ash- D’Amato, and P. J. Palik. 2014. Water table response feeding Lepidoptera. News of the Lepidopterists’ to harvesting and simulated emerald ash borer Society 49: 10–11. mortality in black ash wetlands in Minnesota, Wagner, D. L. 2012. Moth decline in the Northeastern USA. Canadian Journal of Forestry 44: 961–968. United States. News of the Lepidopterists’ Society 54: DOI (10.1139/cjfr-2014-0111). 52–56. Sobek-Swant, S., D. A. Kluza, K. Cuddington, and D. B. Wagner, D. L., D. C. Ferguson, T. L. McCabe, and Lyons. 2012. Potential distribution of emerald ash R. C. Reardon. 2002. Geometroid Caterpillars borer: What can we learn from ecological niche of Northeastern and Appalachian Forests. USFS models using Maxent and GARP? Forest Ecology Technology Transfer Bulletin, FHTET-2001-10. and Management 281: 23–31. DOI (10.1016/j. United States Department of Agriculture, Forest foreco.2012.06.017). Service, Morgantown, West Virginia, USA. 239 pp. Solomon, J. D. 1995. Guide to Insect Borers in North Wagner, D. L., D. F. Schweitzer, J. B. Sullivan, and R. C. American Broadleaf Trees and Shrubs. Agriculture Reardon. 2011. Owlet Caterpillars of Eastern North Handbook 706. United States Department of America. Princeton University Press, Princeton, Agriculture, Forest Service Washington, D.C., USA New Jersey, USA. 576 pp. 735 pp. Wheeler, A. G. 2002. Leptoypha elliptica mcatee Tallamy, D. W. and K. J. Shropshire. 2009. Ranking and L. ilicis drake (Hemiptera: Tingidae): New lepidopteran use of native versus introduced distribution records of seldom-collected lace plants. Conservation Biology 23: 941–947. bugs, with clarification of host-plant relationships. Tanis, S. R., and D. G. McCullough. 2012. Differential Proceedings of the Entomological Society of persistence of blue ash and white ash following Washington 104: 687–691. emerald ash borer invasion. Canadian Journal of Williams, D. J. 2007. Biology of the spiny ash Forest Research 42: 1542–1550. sawfly, Eupareophora parca (Hymenoptera: Tardif, J. and Y. Bergeron. 1992. Ecological analysis Tenthredinidae: Blennocampinae), in Edmonton of black ash (Fraxinus nigra) stands on the shore Alberta. The Canadian Entomologist 139: 269–277. of Lake Duparquet, northern Quebec. Canadian Wilson, S. W., and J. E. McPherson. 1981. Life histories Journal of Botany 70: 2294–2302. of Anormenis septentrionalis, Metcalfa pruinosa, TTD (Tri-trophic Database). 2014. Thematic collection and Ormenoides venusta with descriptions of network: Hemiptera. Complete URL (http://tcn. immature stages. Annals of the Entomological amnh.org/). Society of America 74: 299–311. Tuttle, J. P. 2007. The Hawk Moths of North America: Wood, S. L. 1982. The Bark and Ambrosia Beetles A Natural History Study of the Sphingidae of of North and Central America (Coleoptera: the United States and Canada. The Wedge Scolytidae), a Taxonomic Monograph. Great Entomological Research Foundation, Washington, Basin Naturalist Memoirs, No. 6, Brigham Young D.C., USA 253 pp. University, Provo, Utah, USA. 1356 pp. Usinger, R. L. 1945. Biology and control of ash plant Zacharczenko, B.V., D. L. Wagner, and M. J. Hatfield. bugs in California. Journal of Economic Entomology 2014. A new cryptic Sympistis from eastern North 38: 585–591. America revealed by novel larval phenotype and Vines, R. A. 1984. Trees, Shrubs, and Woody Vines of host plant association (Lepidoptera, Noctuidae, the Southwest, Fifth Edition. University of Texas Oncocnemidinae). ZooKeys 379: 93–107. DOI Press, Austin, Texas, USA. 1104 pp. (10.3897/zookeys.379.5765). Wagner, D. L. 2005. Caterpillars of Eastern North America: A Guide to Identification and Natural History. Princeton University Press, Princeton,

BIOLOGY AND CONTROL OF EMERALD ASH BORER 55 CHAPTER 2: ECOLOGICAL IMPACTS OF EMERALD ASH BORER Area of Expertise/Comments of Area retired wildlife biologist; specialist on game species game specialist on wildlife biologist; retired “Bring­ Ecology; of Wildlife and author Entomology of Professor Home” Nature ing Preserves Nature of Division Ecologist; Region Northeast Preserves Nature of Division Botanist; Conservation Science of Director ecologist Sciences; Biological forest of Professor entomologist Entomology; forest of Professor plant Herbarium; Michigan of University of Director Assistant ecologist plant systematist; (50+ preserves) Areas Natural of Botany; of Coordinator Curator Certifiedbotanist fieldand arborist Ecologist Restoration and Plant ecologist Chief (ODNR); Botanist wildlife biologist non-game Ecologist Wetland Conservation Specialist botanist Scientist; Research species biologist invasive entomologist; Scientist; Research herpetologist Botanist; Sciences; specialist tree Environmental School of and Natural of the Department of Chair Biology and of Professor Sciences; entomologist Physical Affi liation Affi (DEEP) (DEEP) Delaware of University (IND­ Resources Natural of Department Indiana NR) (IND­ Resources Natural of Department Indiana NR) Conservancy (TNC) Nature e Th University State Wayne University State Michigan Michigan of University History Natural of Museum Cleveland Experts Bartlett Tree District Park Metropolitan Cleveland (ODNR) Resources Natural of Department Ohio Wildlife of Division Ohio ODNR- District Park Metropolitan Cleveland History Natural of Museum Cleveland Agri-food Canada and Agriculture Agency Inspection Food Canadian Centre Information Heritage Natural Ontario (NHIC) Guelph of University College Olivet CT-Department of Environmental Protection Protection Environmental of CT-Department Contact May, Dale May, Doug Tallamy, Richard Dunbar, Mike Homoya, John Shuey, Dan Kashian, Deb­ McCullough, orah Reznicek, Anton Bissell, James Boone, Daniel Constance Hausman, Richard Gardener, James McCormac, John Reinier, Semroc, Judy Paul Catling, Bruce Gill, Michael Oldham, Gary Waldron, John Wilterding, State Connecticut Connecticut Delaware Indiana Michigan Ohio Ontario Wisconsin Contacted forest managers, plant ecologists, land managers, and wildlife biologists and wildlife land managers, ecologists, plant managers, forest APPENDIX 1. Contacted

BIOLOGY AND CONTROL OF EMERALD ASH BORER 56 CHAPTER 2: ECOLOGICAL IMPACTS OF EMERALD ASH BORER Position/Area of Expertise of Position/Area Entomologist Research Entomologist Research Entomologist & Project Senior Research Entomologist Leader emeritus Entomologist Research fauna expert western on Entomologist Research & Project Senior Research Entomologist Leader Collections Manager Entomology Entomologist Research Scientist Research emeritus Professor emeritus Professor & Systematics Insect of Professor Associate CollectionResearch Director Entomologist Research Entomologist Research Cook Collection, Arthropod Research Michi- Affi liation Affi Agriculture USDA Lab, Entomology Systematic ServiceResearch (ARS) Agriculture USDA Lab, Entomology Systematic ServiceResearch (ARS) Agricul- of Department Oregon Museum, Insect Survey and ture Lead Taxonomic Curator; Museum Monsanto Company Program; & Management Prevention Pest Insect Agriculture of Department Oregon N/A of Museum National Lab, Entomology Systematic Institution Smithsonian History, Natural Monsanto Company Brigham L. Bean Science Life Museum, Monte University Young Collection Insect A&M University Texas of Museum National Entomology, of Department Institution Smithsonian History, Natural Curator Associate A&M Univer- Texas Entomology, of Department sity Pathology, Plant and Entomology of Department University Auburn A.J. University State gan of Museum National Entomology, of Department Institution Smithsonian History, Natural Wescott, Rick Rick Wescott, Steven Lingafelter, Ted MacRae, Ed Riley, Charles Staines, Patrick Sullivan, omas AZVista, Haden SierraCollection, e Th Atkinson, Th Austin Texas, of University Insect R.Collection, Horace Burke, Manager Collections Curator Executive Wayne Clark, Anthony Cognato, W. Charles O’Brien, Arizona of University Charles Staines, Family Family Eriophyidae Authority Taxonomic Ron Ochua, Tetranychidae Ron Ochua, Hispinae Taxon Acari Acari Coleoptera Brentidae Coleoptera Buprestidae James LaBonte, Ted MacRae, Coleoptera Cerambycidae Ron Alten, Coleoptera Chrysomelidae Shawn Clark, Coleoptera Curculionidae Robert Anderson, Nature of Museum Canadian Coleoptera Contacted taxonomic authorities and principal experts taxonomic arthropodAPPENDIX 2. Contacted for records.

BIOLOGY AND CONTROL OF EMERALD ASH BORER 57 CHAPTER 2: ECOLOGICAL IMPACTS OF EMERALD ASH BORER Position/Area of Expertise of Position/Area Entomology of Professor Collection Curator, Emeritus Professor fauna expert Texas on Entomologist Research Systematist Molecular orator Associate Scientific orator Biology of Professor Entomologist Research Biology of Professor Biosystematist Senior Insect systematist Biology of Professor ffi liate of Texas A&M, University of Texas at Texas of University A&M, Texas of liate ffi Affi liation Affi of Collections, University Research Systematic Museum State Nebraska Collection Arthropods, of California State (CDFA) Biology, Ecology Evolutionary of Department and (UCLA) California Los Angeles of University a Austin Research, Hardwoods Bottomland for Center Service Forest USDA Agriculture USDA Lab, Entomology Systematic ServiceResearch (ARS) of Museum National Lab, Entomology Systematic Collab- current Entomologist, Research retired Institution Smithsonian History, Natural Agriculture USDA Lab, Entomology Systematic ServiceResearch (ARS) Collab- current Entomologist, Research retired of Museum National Lab, Entomology Systematic Institution Smithsonian History, Natural Agriculture and Food of California Department (CDFA)

Sullivan, Patrick Patrick Sullivan, Bill Warner, AZVista, Haden SierraCollection, e Th Agriculture and Food of California Department Curator Executive Quinn, Mike Associate Research Nathan , Schiff Gagné, Raymond Douglass Miller, Dohlen, Carol von University State Utah Gillian Watson, Stephen Wilson, Central Missouri of University Family Family Authority Taxonomic

Taxon Coleoptera Scarabaeidae Brett e, Ratcliff Coleoptera various Henry Hespenheide, Diptera Diptera Agromyzidae Jean Sonja er, Scheff Cecidomyiidae Hemiptera Hemiptera Alyerodidae Aphididae Debra Creel, Roger Blackman, ServiceHemiptera Research Agriculture (ARS) USDA London Museum, History Natural Cicadellidae Hemiptera Specialist Museum Coccoidea Stuart McKamey, Hemiptera Benjamin Normark, Derbidae Amherst Massachusetts, of University Lois O’Brien, Arizona of University (continued) APPENDIX 2. (continued)

BIOLOGY AND CONTROL OF EMERALD ASH BORER 58 CHAPTER 2: ECOLOGICAL IMPACTS OF EMERALD ASH BORER Position/Area of Expertise of Position/Area Entomologist Research Entomologist Research Entomology of Professor Adjunct Entomology of Professor Entomologist Research orator Entomologist Research Entomologist Taxonomic Entomology of Professor Adjunct Wild- and Entomology of Professor Associate Ecologylife orator Entomologist Research moths western of histories life on authority expert moth Arizona Curator Environmental of Professor Distinguished Forestry Sciences and collection manager retired Affi liation Affi Eastern CerealOilseed Centre, and Research Canada Agri-Food and Agriculture of Museum National Lab, Entomology Systemic Institution Smithsonian History, Natural University State Dakota North USDA Center, Research Agriculture Subtropical Service Research Agriculture (ARS) Agriculture USDA Lab, Entomology Systematic ServiceResearch (ARS) Collab- current Entomologist, Research retired of Museum National Lab, Entomology Systemic Institution Smithsonian History, Natural Virginia West Laboratory, Identification Pest Agriculture of Department of Museum National Lab, Entomology Systemic Institution Smithsonian History, Natural Collab- current Entomologist, Research retired of Museum National Lab, Entomology Systemic Institution Smithsonian History, Natural N/A N/A Smithso- History, Natural of Museum National nian Institution of Sciences; University Natural of Institute Follett liate Connecticut affi Henry, Th omas omas Th Henry, Al Wheeler, omas, Donald Th University Clemson Laura Miller, Al Wheeler, University Clemson Noel McFarland, Rand, Evan Terry Fitzgerald, Cortland at (SUNY) York New of University State Family Family Authority Taxonomic Taxon Hemiptera Miridae Michael Schwartz, Hemiptera Pentatomidae Dave Rider, Hemiptera Psyllidae Hemiptera Tingidae Douglass Miller, omas Th Henry, Hemiptera various Hymenoptera Tenthredinidae David Smith, Bartlett, Charles Hymenoptera Eurytomidae Delaware of University Michael Gates, Lepidoptera Geometridae Lepidoptera Gracillariidae Don Davis, Lepidoptera Noctuidae Eric L. Quinter, (continued) APPENDIX 2. (continued)

BIOLOGY AND CONTROL OF EMERALD ASH BORER 59 CHAPTER 2: ECOLOGICAL IMPACTS OF EMERALD ASH BORER Position/Area of Expertise of Position/Area Curator Lepidoptera Entomologist, Taxonomic Scientist Research Inspection Food Canadian with systematist Agency expert taxonomic Biosystematist Senior Insect Entomologist Taxonomic Emeritus Director Fellow Post-Doctoral expert taxonomic Biology Ecology Evolutionary of and Professor expert taxonomic expert taxonomic Fellow Research Honorary Affi liation Affi Collection Arthropods, Florida of State Florida Agriculture of Department Collection Canadi- Insects of National Canadian Ottawa Agency, Inspection Food an California Depart- Center, Diagnostics Pest Plant Agriculture and Food of ment Collection Arach- Insects, of National Canadian Nematodes and nids Cali- of University Biodiversity, of Museum Essig Berkleyfornia- of Museum National Entomology, of Department Institution Smithsonian History, Natural N/A Califor- of University Entomology, of Department Riverside nia at Scientific Sciences,Ecosystem Commonwealth (CSIRO) Organisation Research Industrial and Tuttle, James P. P. James Tuttle, DeBenedictis, John N/A California Davis of University Bohart Museum, Marc Epstein, Fellow Research Jean-Francois Landry, Jerry Powell, Sohn, Jay Family Family Authority Taxonomic Yponomeutidae Taxon Lepidoptera Pyralidae Lepidoptera Sesiidae James Hayden, Lepidoptera Sphingidae Jason Hansen, Lepidoptera Chris Schmidt, University State Michigan Lepidoptera various Lepidoptera various Leafminers Peter Jump, Gall-and David Wagner, Formers various ysanopteraTh various Connecticut of University ysanopteraTh various Charley Eiseman, Debra Creel, ysanopteraTh various N/A Mark Hoddle, Research Service Agriculture (ARS) USDA Laurence Mound, Museum Specialist (continued) APPENDIX 2. (continued)

BIOLOGY AND CONTROL OF EMERALD ASH BORER 60 CHAPTER 2: ECOLOGICAL IMPACTS OF EMERALD ASH BORER

(INRA,

; larvae feed ; recorded on on ; recorded Poaceae Poaceae (Ed. Riley in litt.). Riley(Ed. in litt.).

; mite predator predator ; mite

and

Comment(s) Comment(s) fraxini Aceria name: Valid (Baker 1996) et al., include hosts Polyphagous; Gossyp- Carya, Fraxinus, 2014) Ischyropalpus name: Valid ium, nitidulus 1977) (Landwer, A pion name: Valid cribricolle , deer deer weed, vetch Olea and wild buckwheat 1993) (Bright, Linsleya name: Valid sphaericollis eggs, grasshopper on , lilac Fraxinus on adults (Church suckle honey and 1977; Pinto Gerber, and Bologna,and 1999) American North No known are cerambycids to be ash specialists (Ted be specialiststo ash (Ted in litt.) McCrae Larvae Campsis on feeds radicans but oligophagous Adults is a preferred a preferred is Fraxinus po­ beetlehost; population EAB by impacted tentially Clark in (Shawn litt.) U.S.? U.S.? - to Native vore? Herbi

Invalid Invalid Invalid an Not Invalid Invalid Invalid Taxonomy Taxonomy Taxonomy Taxonomy Herbivore Taxonomy Taxonomy Taxonomy Taxonomy Taxonomically Taxonomically

Fraxinus Fraxinus Fraxinus Fraxinus Fraxinus Fraxinus Fraxinus Specialist Specialist Specialist Specialist Specialist Specialist Specialist Specialist Specialist Specialist Specialist? Valid? Misattributed Misattributed Misattributed Misattributed Misattributed Misattributed

Further Further Research Research Required Required Specialist Not Imperiled Imperiled Not Not Imperiled Imperiled Not

High High High High to to High High to to High Ranking Ranking Previous Current Moderate Moderate Imperiled Not Moderate Moderate Moderate Moderate Imperiled Not Moderate Moderate Imperiled Not Moderate Moderate

Gahan

Species Garm. Garm. fraxini Eriophyes homorus Tetranychus nitidulus Anthicus Pritchard & Baker LeConte Say sphaericollis Lytta Apion porosicolle Apion Gemm. Obrium rufulum Octotoma plicatula (Fabricius) (Fabricius) Taxon Coleoptera Acari Acari Coleoptera Coleoptera Coleoptera Coleoptera Coleoptera Coleoptera Coleoptera Misattributions from Published Literature and Online Sources Literature APPENDIX 3. Misattributions Published from

BIOLOGY AND CONTROL OF EMERALD ASH BORER 61 CHAPTER 2: ECOLOGICAL IMPACTS OF EMERALD ASH BORER

- -

and ; generalist, ; generalist, ; contamination ; contamination ; mycophagous ; mycophagous F. acuminata F. F. acuminata F. (Raymond Gagné Gagné (Raymond

(Drake and Ruhoff , Ruhoff and (Drake and and Comment(s) Comment(s) Porricondyla name: Valid temeritatis in litt.) Gagné (Raymond Contarinia name: Valid canadensis record (Raymond Gagné, Gagné, (Raymond record 1989; in litt.) Solomon from Record D. to (1993) refers et al. apicata; in litt.). Neolasioptera name: Valid fraxinifolia contaminant (Raymond (Raymond contaminant 1989; in litt.) Gagné, (Ray- Generalist predator mond Gagné in litt.) in litt.) Gagné mond general- a definitely “Very saprophage” or phyto- ist in litt.) Gagné (Raymond a wide variety on “Feeds herbaceous woody and of plants” (Wilson and and (Wilson plants” 1981) McPherson, ligu Forestiera are Hosts strina 2002) (Wheeler, ligu Forestiera are Hosts 2002) (Wheeler, Osmanthus are Hosts strina Lyonia 1965) U.S.? U.S.? - to Native vore? Not an an Not Herbi Herbivore

Invalid Invalid an Not Invalid Invalid Invalid Invalid Taxonomy Taxonomy Herbivore Taxonomy Taxonomy Taxonomy Taxonomy Taxonomy Taxonomically Taxonomically

Fraxinus Fraxinus Fraxinus Fraxinus Fraxinus Fraxinus Fraxinus Fraxinus Fraxinus Fraxinus Fraxinus Fraxinus Fraxinus Fraxinus Specialist Specialist Specialist Specialist Specialist Specialist Specialist Specialist Specialist Specialist Specialist Specialist Specialist Specialist Specialist Specialist Specialist? Valid? Misattributed Misattributed Misattributed Misattributed Misattributed Misattributed Misattributed Misattributed Misattributed

Not Imperiled Imperiled Not Not Imperiled Imperiled Not Not Imperiled Imperiled Not Not Imperiled Imperiled Not Not Imperiled Imperiled Not Not Imperiled Imperiled Not Not Imperiled Imperiled Not

High High High High High High High High- High- High to to High Ranking Ranking Previous Current Moderate Moderate Imperiled Not Moderate Moderate Moderate Moderate Imperiled Not

-

(Spinola) (Spinola)

Species Felt Colpodia temeritatis Continaria canadensis Felt Dasineura spp. Dasineura ora fraxinifl Lasioptera Felt Lestodiplosis fraxinifolia Felt Rhizomyia fraxinifolia fraxinifolia Rhizomyia Felt Anormenis septentrion alis Leptoypha elliptica McAtee Drake ilicis Leptoypha Leptoypha mcateei Drake Drake Taxon Diptera Diptera Diptera Diptera Diptera Diptera Diptera Diptera Diptera Diptera Hemiptera Hemiptera Hemiptera Hemiptera Hemiptera Hemiptera Hemiptera Hemiptera APPENDIX 3. (continued) APPENDIX 3. (continued)

BIOLOGY AND CONTROL OF EMERALD ASH BORER 62 CHAPTER 2: ECOLOGICAL IMPACTS OF EMERALD ASH BORER

Comment(s) Comment(s) Leptoypha name: Valid minor species (Hod- European 1988) kinson, Lignyodes name: Valid bischoffi Lignyodes name: Valid helvolus & reared bulk Wasps ash; with associated only species likely parasitoid parasitoid species likely 2007) (Tuttle, Podesia of form Color oligophagous syringa; (Keen, 1958; David Smith Smith 1958; David (Keen, in litt.) Jamaica from recorded Tethida barda barda Tethida name: Valid species Neotropical Duckworth, and (Eichlin 1988) U.S.? U.S.? Species troduced troduced Extralimi- tal Species - In Exotic/ - to Native vore? Not an an Not Herbi Herbivore

Invalid Invalid Invalid Invalid Invalid Invalid Invalid Invalid Invalid Invalid Taxonomy Taxonomy Taxonomy Taxonomy Taxonomy Taxonomy Taxonomy Taxonomy Taxonomy Taxonomy Taxonomically Taxonomically

Fraxinus Fraxinus Fraxinus Specialist Specialist Specialist Specialist Specialist? Valid? Misattributed Misattributed

High to to High High to to High Moderate Moderate Moderate Moderate Not Imperiled Imperiled Not Not Imperiled Imperiled Not Not Imperiled Imperiled Not

High High High High High High High to to High to High High to to High Ranking Ranking Previous Current Moderate Moderate Moderate Moderate Moderate Imperiled Not Moderate Moderate

spp.

Species Leptoypha nubilis Drake Drake Psyllopsis fraxinicola fraxinicola Psyllopsis Forst. bischoffi ysanocnemis Th Blatchley Th ysanocnemis helvola helvola ysanocnemis Th LeConte Eurytoma Podosesia fraxini (Lug- fraxini Podosesia Tethida cordigera cordigera Tethida (Drury) ger) ger) Manduca brontes Manduca - - Taxon Hemiptera Hemiptera Hemiptera Hemiptera Hemiptera Hemiptera Hemiptera Hymenop tera tera Lepidoptera Hymenop Lepidoptera tera tera (Beauvois) APPENDIX 3. (continued) APPENDIX 3. (continued)

BIOLOGY AND CONTROL OF EMERALD ASH BORER 63