·et-t fua-~: Ash in Pennsylvania

PROGRESS and

FUTURE DIRECTIONS in Research on the Emerald Ash Borer by Therese M. Poland, Research Entomologist, Ecology and Management of Invasive Species and Forest Ecosystems, USDA Forest Service

When the emerald ash borer (EAB) was discovered near Detroit, Michigan in July 2002, very little was known about it

other than the fact that it was killing large numbers of ash trees throughout a widespread area in southeast Michigan

(Poland and McCullough 2006). Ash mortality in the area had been noted for a few years, but was attributed to ash

decline until damage and symptoms including galleries and exit holes became so prevalent that it was clear the beetle

damage was not secondary but was in fact the causative agent of mortality. The beetle was identified by Eduard Jendek

as Agrilus planipennis Fairmaire (Coleoptera: Buprestidae), an exotic wood boring beetle native to Asia (Haack et al. 2002).

Some basic aspects of its biology and general descriptions were available and translated from Chinese textbooks (Chinese

Academy of Science 1986, Yu 1992), but nothing was known about how to detect or control it. Hence, research scientists

quickly initiated projects to learn about this invasive species and develop tools to manage it .

....u.-,_.,. ... f-11-1 •• ""'kttf ~• ..... lifJlilwr. JW.w/IMi/kr, USDA Forat

BIOLOGY Research on the biology of EAB in

North America demonstrates that it is

generally quite similar to the life cycle

described for EAB in China (Chinese

Academy of Science 1986, Yu 1992).

The life cycle is typically completed in

one year, but two years may be required,

especially in vigorous hosts with light

infestations, in cooler climates, or when

eggs are deposi red late in the season

20

(Cappaert et al. 2005, Wei et al. 2007,

Wang et al. 2010, Tluczek et al. 2011 ).

Two years may also be required to

complete development in cut logs or

firewood, particularly when the wood

has dried (Petrice and Haack 2007) .

Adult beetles chew their way out of

trees leaving 0-shaped emergence holes.

In the Great Lakes region of North

America and at similar latitudes in Asia,

adults begin emergence in May or June,

activity peaks in late June to early July,

and the flight period usually ends by

September (Cappaert et al., 2005; Wei

et al., 2007; Wang et al., 20 10). Adults

are most active on sunny days when

air temperatures exceed 25°C [77°F]

(Wang et al., 20 IO) and often rest in

bark crevices and on leaves during rainy

or cool weather (Rodriguez-Saona et al. ,

2007). EAB adults feed o n ash leaves

SUMMER 2014

-

PROGRESS AND FUTURE DIRECTIONS IN RESEARCH ON THE EMERALD ASH BORER

(Fig. l) throughouc cheir life which

can lase for several weeks (Bauer et al.

2004, Lyons ec al. 2004, Wang et al.

20 IO). Adulcs begin macing afcer 5-7

days of macuracion feeding and females

feed for an addicional 5-7 days before

beginning ro deposic eggs in bark cracks

or crevices.

Eggs hacch within cwo weeks and

the larvae feed in che phloem and

cambium through aucumn creating

serpentine-shaped galleries thac are

packed wich frass. Larvae pass through

four developmental inscars (Cappaert

ec al. 2005) and most complece feeding

in October or November. Pre-pupae

overwinter in cells about l.27 cm deep

in che sapwood of thin-barked trees

or in che outer bark of chick-barked

trees. Pupation begins in mid-April and

continues into May, followed by adult

emergence approximacely 3 weeks lacer.

Some EAB overwinter as young larvae in

their galleries and chen require a second

year of development before emerging as

adults (Cappaert et al. 2005, Tluczek et

al. 2011). EAB prepupae are intoleranc

of freezing and survive winter by

accumulacing high concentrations of

glycerol and other antifreeze compounds

to achieve supercooling points of abouc

-30°C [-22°F] (Croschwaice et al. 20 l l).

On-going research is exploring many

ocher aspects of EAB biology and

ecology such as overwinter survival,

temperature influences on larval

development, nutritional requirements

for development of artificial diet and

rearing methods, and factors chac

influence macing and reproduction.

HOST RELATIONS Ash is the only larval hose reported for

EAB in China {Yu 1992; Liu et al.,

2003; Zhao ec al., 2005). EAB was

synonymized [equated with] wich cwo

ocher Asian Agrilus species and one

subspecies: Agrilus feretrius Obenberger

(type Taiwan), Agrilus marcopoli

Obenberger (type Mongolia), and

Agrilus marcopoli ulmi Kurosawa (type

Japan) Oendek and Grebennikov, 20 11).

Ocher tree genera (Jug/ans, Pterocarya,

PENNSYLVANIA FORESTS

and Ulmus) have been reported as

larval hoses in Korea and Japan for

A. marcopoli and A. marcopoli ulmi

(Ko, 1969, Akiyama and Ohmomo,

1997). There are 27 species and one

sub-species of ash, Fraxinus, native

to Asia (Wei 1992) of which emerald

ash borer mainly infests Manchurian

ash (F manchurica), Korean ash (F

rhynchophylla), and C hinese ash (F

chinensis) (Yu 1992), generally attacking

only stressed trees. le also attacks the

North American species, green ash (F

pennsylvanica), white ash (F americana),

and velvet ash (F velutina), which are

commonly planted in China (Liu et

al. 2003, Zhao et al. 2005). North

American ash species are attacked at

higher levels than native Asian ash

species in China; at one site 95% of

orch American green ash trees were

moderately infested while no infestacion

was found in Korean ash trees of similar

size planted side by side (Liu et al.

2007). In its introduced range in North

America, emerald ash borer is able co

attack every species of ash it has so-far

encountered including black (F nigra),

blue (F quadrangulata), green, pumpkin

(F profimda), and white ash and even

healthy trees are attacked and killed

(Poland and McCullough 2006).

Anulewicz et al. (2008) compared

adulc landing and oviposicion on

logs of several North American ash

species and on non-ash species chac

are congeners [belonging co the

same genus] of hoses reported for

che synonomized Agrilus species in

Korea and Japan. Non-hoses included

American elm (Ulmus americana),

hackberry ( Celtis occidentalis), black

walnut (Jug/ans nigra), shagbark hickory

( Carya ovatae), and Japanese tree li lac

(Syringa reticulatae). Adulcs landed and

oviposiced less frequencly on non-ash

logs compared co ash logs and no larvae

were able to survive, grow, or develop

in non-ash logs. Although all species of

North American ash appear susceptible

co emerald ash borer, preference and

susceptibility vary among ash species.

For example, Anulewicz et al. (2007)

Forest Set·vice tech11icz'ans checking a double decker trap for emerald ash borers (photographer, Therese Pola11d, USDA Forest Se1·vice)

found signiflcancly higher canopy

dieback and emerald ash borer attack

density in green ash than in white ash

trees at che same sices, and in white

ash compared to blue ash crees ac the

same sites. Similarly, Tanis et al. (20 12)

found greater persistence of blue ash

compared co white ash and suggested

ic was more resiscant co emerald ash

borer. Black ash experienced greater

£AB-induced mortality chan white or

green ash in 31 permanent vegetation

plots in forest stands in Upper

Huron River Watershed, although all

three species were severely impacted

(Gandhi ec al. 2008). Differences in

suscepcibility co emerald ash borer

among ash species have been found co

be related co differences in host volatiles,

nucricion, and defense compounds.

Pureswaran and Poland (2009a) found

thac emerald ash borer adults preferred

co feed on green, white and black

compared co European (F. excelsior),

blue, or Manchurian ash and che relacive

amounts of anrennally-accive volaciles

varied among ash species. Emerald ash

borer also prefers co feed on mature

leaves compared to newly flushed leaves,

on leaves grown in sun compared co

leaves grown in shade, and on leaves

from trees chat had been scressed by

girdling compared co leaves from

21

PROGRESS AND FUTURE DIRECTIONS IN RESEARCH ON THE EMERALD ASH BORER

Injecting ash tree UJith systemic iusecticides (photographer, Therese Poland, USDA Fo1'f!st Service)

healthy trees (Chen and Poland 2009).

Preferences were related ro lower defense

compounds (trypsin and chymorrypsin

inhibitors) and greater nutritional value

with higher levels of amino acids and

protein ro carbohydrate ratios.

In a common garden study near rhe

initial infestation area in Michigan,

emerald ash borer arrack densities and

rree mortality were significantly higher

in white and green ash culrivars rhan

in a Manchurian ash culrivar (Rebek er

al. 2008). Ir is hypothesized rhar rhe

greater susceptibility of Norrh American

ash species compared ro Asian ash

species ar field sires in borh China and

orrh America may be due ro resistance

mechanisms char developed in Asian ash

species rhrough rheir close evolutionary

history with emerald ash borer rhar is

lacking wirh Norrh American species.

Eyles er al. (2007) analyzed phloem

chemistry of different ash species

and found differences in rhe array

of phenolic compounds among ash

species and rhar hydroxycoumarins and

calceolariosides A and B were unique

ro Manchurian ash. More recendy,

Whitehill er al. (2011) also found

differences in qualitative phenolic

profiles among ash species wirh

Manchurian ash being mosr differenr

from rhe green ash variety 'Parmore',

and white ash variety 'Autumn Purple'

culrivars. However, they found rhe

hydroll.J'COumarins and calceolariosides

were present in highly susceptible black

ash and European ash, refuting their

role in resistance ro emerald ash borer.

Similarly, Cipollini er al. (201 1) found

22

differences in phenolic profiles among

ash species, with 9 phenolics unique ro

Manchurian ash. Different species of

ash also respond differently ro emerald

ash borer adult and larval feeding. In

response ro adulr feeding green and

white ash had higher levels of induced

volarile emission rhan black ash, levels

of roral phenolics decreased in whire

ash, and chymorrypsin inhibitors were

increased in black ash (Chen er al.

20 lla). Larval feeding also induced

volatile emission of (E)-~-ocimine and

(Z,E)-a-farnesene in black ash, increased

levels of carbohydrates and phenolics,

and decreased levels of proreins and

amino acids (Chen er al. 2011 b).

Emerald ash borer larvae utilized green

ash amino acids more efficiently rhan

those of whire or black ash and was were

able ro eliminate phenolics through

excretion and enzymatic conversion

(Chen er al. 2012).

Asian ash species are being explored

as a porenrial source of generic

resistance and efforrs ro breed EAB

resistant ash are ongoing (Koch er

al. 20 12). A small proporrion of ash

have survived and remain healthy

in ash stands throughout Michigan

and Ohio where over 99% of rhe ash

trees have been killed by EAB. These

lingering ash trees are being evaluated

as a potential source of resistance genes

in native ash populations (Knight er

al. 2013). Genomic sequencing of

Asian and Norrh American ash species

has also been conducted ro provide

a molecular foundation for targeting

resistance breeding (Bai er al. 2011),

and transcripromic studies of EAB are

exploring mechanisms by which larvae

detoxify host defenses (Mirrapalli er al.

2010, Rajarapu er al. 2011 ,).

SURVEY AND DETECTION Initial delimirarion of the infested

area in Michigan was based on visual

surveys using external symptoms. Visual

surveys along with tracebacks of nursery

srock movement our of Detroit were

used ro derecr new infesrarions of EAB

through 2003. Ir became apparent

that visual surveys were nor effective

for derecring low density infestations of

EAB. Research demonstrated rhar adult

beetles are amacred ro volatiles emitted

by stressed ash (Rodriguez-Sanoa er al.

2006) and preferentially oviposir on

girdled trees (McCullough er al. 2009a,

2009b). As a result, the Michigan

Department of Agriculture implemented

a statewide grid of girdled rrap trees

in their EAB survey in 2004. Girdled

trap trees were visually inspected during

the summer ro collect any captured

EAB adults from sticky bands. Trees

were felled during the fall/winter and

bark was peeled from sections of rhe

upper trunk and canopy ro locate any

EAB larvae and galleries (Hunt 2007,

Rauscher 2006). Trap trees were used for

detection surveys in Michigan and several

surrounding states through 2008. While

girdled trees are the most effective rool

for EAB detection (McCullough er al.

2011, Mercader et al. 2013), debarking

trees to locate larval galleries can be labor

intensive and costly. Moreover, suitable

rrees may nor always be available or

accessible, especially in urban areas or

when mulri-year surveys are needed.

Considerable research has, therefore,

been invested in efforts to develop

effective artificial traps and lures

ro attract and capture EAB adults.

Artificial traps were developed

incorporating arrracrive olfactory and

visual cues and first implemented for

derecrion surveys in 2008 (Crook and

Mamo 20 IO). Volatiles from ash trees

leaves and bark elicit antennal responses

and are arrracrive ro EAB (Rodriguez­

Saona er al. 2006; deGroor er al. 2008;

C rook er al. 2008; Grant er al. 2010,

2011; Poland er al. 2011). Many of

rhe volatiles present in ash bark are also

present in Manuka oil (Crook er al.

2008).

The beetles are also amacred ro specific

wavelengths of violet and green lighr

(Crook er al. 2012), and females are also

sensitive ro red ranges of rhe spectrum.

EAB adults are attracted ro traps colored

different shades of green or purple hung

SUMMER 2014

PROGRESS AND FUTURE DIRECTIONS IN RESEARCH ON THE EMERALD ASH BORER

in che open or in che canopy of ash crees

(Francese ec al. 2005, Crook ec al. 2009,

Francese ec al. 20 10). Males, chac cend

co hover near the canopy of ash trees, are

captured in higher proporcions in green

craps hung in che canopy of ash trees;

whereas, females, chac oviposic on che

crunks of ash crees are capcured in higher

proporcions in purple craps hung below

che canopy (Crook and Masero 2011).

In addicion, there is recenc evidence

of close range or conracc pheromones

involved in mace recognicion and

macing behavior (Lelico ec al. 2009;

Pureswaran and Poland 2009b) and a

female-produced volatile pheromone,

cis-laccone, chac increases accraccion of

males co green canopy craps baited with

host volatiles (Silk er al. 2009, 2011;

Ryal! ec al. 2012).

Traps used for detection surveys have

evolved over the years co incorporate

the laresr research findings. Currently,

national detection surveys conducted by

USDA Animal Plane Heal ch l nspeccion

Service (APHIS) employ scicky purple

prism traps hung in che canopy of

ash trees and baiced wich che ash leaf

volatile, cis-3-hexenol, and Manuka

oil (APHIS 2014). Ocher promising

craps under evaluat ion in pilot surveys

include green multiple fu nnel craps

haired with cis-3-hexenol, green sticky

prism craps hung in the canopy of ash

crees baiced wich cis-3-hexenol and rhe

EAB pheromone cis-laccone, and purple

and green double decker craps (Fig. 2)

com posed of a 10 foot PVC pole co

which rwo sticky prisms are arrached

near rhe cop and haired wirh cis-3-

hexenol. Conrinued research on close

range pheromones and improvements to

trap designs and lures will lead co more

effective detection cools.

CHEMICAL CONTROL Soon after EAB was discovered studies

were initiated co evaluate d ifferent

insecticides as a means of rapid control.

Early studies reseed canopy sprays wirh

persistent insecticide formulacions

(e.g., bifenthrin, cyfluthrin). Sprays

were nor popular because of problems

PENNSYLVANIA FORESTS

Tetrastichus plmiipeim isi, larval parasitoid of e111erald ash borer (photograph.,; David CapptJert, D eparh11e11 t of Entomology, Michig1111 State U11iversity)

such as pocential drift, environmental

contamination, non-rargec impacts on

pollinacors and beneficial predacory

inseccs, and possible applicator exposure.

Syscemic inseccicides are applied by

injeccing produces di rectly into the

crunks of crees (Fig. 3) or by applicacion

co che soil or as a basal trunk spray,

eliminacing most problems associaced

wich cover sprays (H erms ec al. 2009,

McCullough ec al. 2011, H erms and

McCullough 2014). However, once

high densicies of EAB larvae have

injured che vascular syscem of an ash cree

cranslocacion and efficacy of syscemic

inseccicides are impaired.

Inicially only a few syscemic inseccicide

produces were available, mainly wich

imidacloprid as che accive ingredienr

and efficacy crials yielded inconsistent

resulcs (Herms er al. 2009) . New

syscemic inseccicides are now available

and applicacion technology has

improved. An emameccin benzoace­

based inseccicide was fi rsc regiscered

in rhe US in 20 I 0 and is currently rhe

mosr effective produce available (H erms

and McCullough 2014), providing rwo

co three years of nearly complere EAB

control (Herms er al. 20 I 0, Smitley

er al. 20 I 0, McCullough er al. 2011,

2012,). Many mu nicipalities and private

landowners are now proceccing ash trees

from EAB wirh emamecrin benzoare.

Dinorefuran is a highly soluble new

generation neonicorinoid produce.

Ir can be applied as a basal trunk spray

and is effective if applied annually.

Ir is popular among arborists when

many small crees require treatment

(McCullough er al. 2011, Herms and

McCullough 2014). lmidacloprid-based

insecticides must be applied annually

and continue co be used for EAB control

wich variable efficacy. Azadirachtin

is a natural insecticide derived from

che neem cree (Azidirachca indica).

Azadirachcin produces are noc coxic to

EAB adulcs, hue impair reproduction

and control young larvae. They provide

one co rwo years of cree proceccion,

depending on local EAB densicy

(McKenzie er al. 2010).

Economic analyses have shown chat

insecticide creacment is highly cosc

effeccive in urban sercings compared

co che coses of removing crees killed by

EAB (Kovacs ec al. 2010). Treating a

porcion of che crees in a given area may

also slow the race of EAB populacion

growth (Mercader ec al. 2011).

H owever, broadscale applicacon of

syscem ic insecticides is noc praccical for

23

PROGRESS AND FUTURE DIRECTIONS IN RESEARCH ON THE EMERALD ASH BORER

EAB management in forests, woodlots,

riparian zones or other natural areas due

to high costs associated with individual

tree treatment and environmental

concerns rhar may limit use of

insecticides in natural areas. Effective

biological control of EAB, whether by

native natural enemies or introduced

parasiroids, may be critical to preventing

the functional loss of many ash species

in foresr ecosystems across Norrh

America (Klooster et al. 2014, Knight

et al. 2013). Biological control and

systemic insecticides are not mutually

exclusive and may result in additive or

even synergistic negative effects on EAB

populations (Barclay 1987, Berec et al.

2007, Suckling et al. 2012). Future

research will focus on new and improved

insecticide formulations and application

methods along with integration of

insecticide control with other tactics for

area-wide management of EAB.

NATURAL ENEMIES AND BIOLOGICAL CONTROL In 2003, surveys were initiated for

natural enemies of EAB in Michigan

and China. In southeast Michigan (2003-2004) and a more recent

survey in Pennsylvania (2008), <4%

of EAB larvae were found parasitized by native and naturalized non-native

parasitic wasps (Bauer et al. 2012) .

Ectoparasitoids are the most common

group attacking EAB larvae, and of

those, Atanycolus spp. and Spnthius spp.

(Braconidae) are the most prevalent

in Michigan. No EAB egg parasitoids

have been found in Norrh America.

In northeast China rhree parasitoid

species were found arracking EAB:

Oobiw agrili (Encyrridae), an egg

parasitoid; and Tetrastichus planipennisi

(Eulophidae) and Spathius agrili

(Braconidae) rwo larval parasitoids

(Fig. 4). At one site in Jilin province,

combined parasitism by 0. agrili and

T plnnipennisi was estimated to reduce

EAB densities by 74% in green ash

trees (Liu et al. 2007). S. agrili was

later found at that site but was rare.

These three EAB parasitoid were studied

extensively for non-target impacts in

24

containment/quarantine laboratories,

and approved by US Environ mental

Protection Agency for environmental

release in late July 2007 (USDA APHIS

2014b)

The three species of Asian parasitoids

were first released at sires in southeast

Michigan in 2007 (USDA APHIS

2010). In 2009, theAPHIS EAB

Biocontrol Facility in Brighton, MI

became operational, after transfer

of improved rearing methods and

parasitoid cultures from the Forest

Service and Center for Plant H ealth

Science and Technology (CPHST)

laboratories. Production has increased

annually, and in 2012, more than

350,000 wasps were released in 14

states. Several releases have resulted

in successful establishment, although

establishment of S. agrili in Michigan

has been limited, possibly because of

cold weather (Duan et al. 2009, 2010,

2012). Exploration and evaluation

of additional Asian parasitoid species

for potential introduction are ongoing

(Duan et al. 2012), including another

Spathius species, S. galinae, native

to Russian Far East, that may be

more adapted to cooler climates

(Belokobylskij et al. 20 12). Current

research is evaluating effects of the

parasitoids on EAB population growth

rates and ash tree health and will require

assessment over multiple years.

FUTURE OUTLOOK FOR ASH MANAGEMENT Survey and management tools

developed by research are now being

implemented in an integrated strategy

and tested in a multi-agency pilot SL.ow

A.sh M.ortaliry (SLAM) study. The

approach incorporates I) Surveys of

EAB infestation and distribution using

arrificial traps; 2) ash host survey to

determine area at risk and plan location

of derecrion traps and treatments;

3) population suppression through

insecticide rreatment of landscape

trees and trees in a buffer zone around

positive detections, clusters of girdled

trap trees that are felled and debarked to

detect and destroy beetles, removal and

utilization of infested trees, and release

of natural enemies for biological control;

4) regulatory control to prevent arrificial

movement; and 5) public outreach. The

SLAM approach is mosr likely to be

successful when implemented in areas

with new infestations that are relatively

small and isolated. Landscape-level

management strategies including SLAM

and biological control, insecticide

treatments in urban areas, collection

and preservation of ash seed, and

development of more resistant ash,

offer hope for the protection of ash and

persistence of the genus at some level in

urban and natural forests. 0

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Chinoc Aadcmr of Sden<e. lruciru« of Zoology 1986. Apil<11 nurcopoli ~- Agricuhut lrucru ofChuu •pan IJ, p. 44;. Chiiu .\gricul""' I'=. Bri1cng, Chiiu.

Cipollm1. D .. Q. Wang. J. G. A. 111mchill. j. R. Po.di. P Bonello. and D . . \. Her'"' 2011. i);,unguishing defense chmctrru<i<.1 in che phloem of ash ~p«ics rcsis1;;im and ~usccpublt 10 cmcrakl .ish borer. JoumJJ of Chemial f.col<>g)· 3":4)0-j9.

SUMMER 2014

1!11!

PROGRESS AND FUTURE DIRECTIONS IN RESEARCH ON THE EMERALD ASH BORER Cruol D. J. •nd I'. C. "'"''°· !010. Chama! mil~ of 1h< ,mmiJ ash bor?T

Arrlusp~mpmu.fau,.../ofO..."'' Eto"rf .l6: 101-11!.

Co'*- D .• A. IJuimun. ). A. frm<oc. I. fu><r. T. \I l'ubnd. A.). 1.i•)tt. and r. C. Mastro. 2008. Dn·dopmrm of a host-based snniochcmial lure for inpping cmmld ash bor?T, AriJ"' ~"pniniJ (Colcopicn: il<lprcsti<b<I. Em1ronmcn1~ Emomolop r- Ji6-361.

Crook. 0. J., Khrimi.in. A .. Cos.st. I. Frmr, ;1.nd V. C. Mrnro. 2012. lnflutn<tof inp oolor •nd 00.1 >"ob1ila on ap1u1< of ih< .mmlJ ash bom (Cclrop<m: Bup1audxl jounul of&ononuc Emomology IOI: 4!'>-ff.

Cro>1h-.i1e ). C .. S. Sobd<. D. B. Lyons. M.A. lkrnard>. •nd B. j. Sm,~ir. 2011. Th< "'"'inicring ph'""°s' of ih< .mmlJ ..Ji bom. Arri• ~.,pninu hi= ICokvpma 8"pm1idxl jounul of ln><tt l'h~'Siolog) 5-, 16&--J

.X GroOI. P.. G. G. Gram. T. M. Pobnd. R. SJmlmh. L Bud.m. R. II'. ~011. L \bdlo1uld. and D. Pm. !008. Elmroph~'iologial "'f'O'U' and mrxtl<ll\

ii .mmlJ ..Ji bom 10 gnm bf ,,,U11lo (GL\'s) nm11cd ~ h<.•1 fulus<. Journal of Chemical Erolog.> 34: W0-1179.

Du...j.j .. R. \I FU<Sltt.). llildongtr. PB. foioi.S. &rth.and \. E. lpi..~igtt. !009. P=i1oid. mxling 1hc cmmld ..Ji bor?T (Cokopitta Bup1oti<b<I m 11ies1nn Pcnnsyk1nU. FloriJ.a Emomologis1 92: 588-92.

D.un. JJ. . .II. D. L'lyJ1m. L \ &un. j. Goold. and R. "'" Dncsdit !010. \teasunng 1hc imp.act of bto1ic factors on popubuons of imnuturr nnmki bh bom (Cokvpma 8"p1<S1idii:I. Enrno(\ffi(nul En1ornolofl .\9: l\IJ.-!!.

Own. J. ) .• L S.utt, K. j. \hdl. and R. G "'" Dno<h<. 201! l'opuhuon mponscs of h)·mroop1rnn p,Jr~i1oids to 1h< emmld uh borer (Colroptm: Buprcsti<bc) in rtemlh im~ .ims in nonh emir.al l'ni1cJ Smo BioConuol 5-· 199-!I~).

hln . .\ .. II. Jones, K. Riedl. D. Gpollrni. S. Sch"'n• K. Chan. 0. A. Hc11n<. and~ Bonello. roo-. U>ml""'"" phlonn chcminn .i\bndiun.m tfr.oonlU 11Ulflls1•1m<itl ind ~v \onh ,\mm.;m i.Ji \pn.10 ~F ll"ff71(Jlfl and F P"''"Jf•,.r<'). Joonul ofChttnial Ec"'°&l Jl: 1430-IHS.

F""'"'· ). A .. D. j. Crook. I. f..,,,. D.R. un«. A. j. '"'"'·and I'. C. \J.suo. !OIO. Opumu.a1"" of"'!"°"" for 1h< tmmld ..Ji bom. A.!"/"'~"!"'"' 1Colrop1m: Buprcs11du). joonul of Economic Eniomok>g)· I O.l: I !35-1241.

G..Jhi. K. J. '-, A. Sm11h. R. P Long. R. .\ .. J. foioi. and D. A. Hmn> !OOl>. 1111«-Yw l'rog=ion of En"1lld Ash Bor?T-lnduccd Dcdin< and .lloruli1y in Sou1heasmn Michigan. p. r. In V. M.isno. 0. Lance, R. Rardon. D. Pm lcompilm). En"1lld Ash llo<tt Rocarth and Tcdinology &dopmcn1 \!ming. LIDA Fomt Scnict. Moipn"''"· 11~1. FHTET-roo~-.

Gram. G. G. K. L. R)>ll. D. B. L~~ru. •nd M. M. Abou·Z.id. 2010. Oilftt<111ul "'I"""' of nuk and l<nuk .mmlJ "h boms !Col .. lluproud"I 10 ZJ-} hac:nol and \bnuka oil. ~ ... nul of Applocd En1omolos-· l}I; !(>.l.l.

Gram. G. G .. T. \1 Pobnd. T. Cuumiuro. D. B. I ~vns. and G C jonn !O 11. WP'"""' of .\bk and Fmuk Emtt~d \sh Bor?T ICokopitt~ il<lpl<Slidx Ropons<1 10 l'hrxbc Oil and IZJ-J-Hmnol l.u1t1 rn I iglu G""' Pn,m fop, jounul ofEconomk Eniomol<>g." 104: l'J·l-9.

Haxl R. .l. E. )tndcl H. I.Ju. K. R. \ludum. T. R. l'nricc. T. \I. Poland. and H. Yr. 2001. Thr tl1l('nld ash bom: A new O:Oll{ po:1 in ;\onh Amrric.i. '\.-•k1<cr of 1h< Michii;Jn En1omolopal Socitty 4' (JMl: 1-1.

Hmiu. D .. l. D. G .lkCullough. D. R. \m"ki• C. E s.dof. R. C 11 illiun""'. and P. L :\1xon. 2009. lnsmKidroptions for pro1tc1ingash trttS from anm.ld bh bo1tt. ~onh Cm1ral IP\! Cm1a 8"lk11n. I! pp Auibhlc "' b11p:I/ .,.,"'""'·ancrM.uhbortr.info, filo. \fulti11.nt_F.AB_lnsa.ucidt_fil't .. 'hm.pJf ("c...cd4)un<l0141.

Hcrm" D. A. !010. Mul1irar t''11u..nie>M of srmmic irumicido for control of tmmld bh bom. pp. 'J.-;. In l'ro=lingi of 1h< Emtt~ .l.h llo<tt R=<th.ndTo;hnoi<>g) Dc.dopmcni .li«1ing. D. l•ncc.J. Buck. D. Bi111on. R. Rrudon. \ ' M"'ro (cdironl.Oa. !0-!1.2009. PnMwgh. PA. l:SD.\foc. Hwdi Tcchnol. En1ttp"" T"'". FHTET-!010-01.

Hc11ns. D. A. and D. G. McCullou,)i. !014. Emor~d >Sh bo"' m•.sion of:\onh America hb<O<y. ~·cc~. 1m!"'-I and nwugcm.01. \nn...l Rt\ic­oiEn1omolop 59: 13- JO.

Hum. L 2007. Emera.Id ash born ~mr updi1r: Ohio, p. 2. In\'. Mis1ro. D. Un.:c. R. Rardon. and G. P.im ICompiknl. Pnxmlingsof die En"1lld .\sh llo<tt and Asian Longhom<d lknk R=rch and T«hnoi<>g) Dc.dopmm1 .lt«ung. Cincinna11. OH. !') Oa. - 2 Nov. 1006. lJ.S. D<p.nmeni of ~1urt. Fo"" Scnicc Publiaaon FHID-200'-04. \loipn1o•m \lY.

jmd.l. E. .. J \'I. Grd>Cn1111.0<. 2011 .1rrl"' ic.lropim. Bup1n1r<bcl ofw1 .Asia.)"' F..bc. P..guc. J6l pp.

Kloo.tt. II I D. A. Hmn> t;. S. Knip11. C. P. H'""' D. G . .lk{;il!oogh. A. \m,.h, K. J. K. Gandhi. •nd J. Cardrna. 1014. Ash (/'nmn., tpp.) monal1~. r~nmtion. ;ind strd bank dnumia in milai hudwood fornl\ fo11cJ.,.ing rm..,ron b. cmmld bh bom cjznU., ,i..,pninu). Biologial IO\bions. /, , ....

'-igh1. K. S .. ). P. B'°"n •nd R. P. Long. 2013. F.aon .ifccnng 1h< su1Yi-..J of ash frmn• spp.1 ""' info1cd b. ttntt.!d ash boner !A_rilas 1Lmpninu). B.ologicall"'"ionsll:n·.lSJ.

PENNSYLVANIA FORESTS

~o. J. H. 1969. A Lisi of Forest lnsm Pot~ m Kor~. Fornt Rrsanh lnsmu1r, Seoul. Korta

Meli. j. L. D. \l. Carty. ~. S. 1'nigh1. T. l'ubnd. D. A. Hmm. \I. E. .\bion. 1011. Br«d1ng ma1rgics for 1he dr•c'°Pmrn1 of rmmld :uh borrr-rcsim.nt '\Olth A.mcrian ash. pp. !35-!39. In R. A. Sn1<Llo. A. D. \""hul J. T. Klitju""- K. .II. r.lmi<n. J. M. Alcsandcr. S. J. fonltl led>.) l'rocmlmp of tht 41h lnmrutionil \\'orbhop on Genni" Hos1·Parui1r lmrr.actions in Foo:stry. C.n. USDA For<>1 Scn·ict l'xifK Soudi•<>t R=rch lmon. Tcth. Rq>. PS\l.(,IR-!40.

K"'·"'· K. F .. R. G. Haigh1. D. G. McCullough. R. J. M<1ad<1. N. \l. iq;m and A. \I. Ud>hold. 2010. Cost of po1<111w nnmld ash bom dam.gt in L'.S. commumuo. !009-2019. EcologJal Economio 69: 569-;-8.

Ldi10. J. P.. K. BOriiczky. T. H. jond, I. Frllt1. V. C. Mamo. J. H. Tumlinson . and T. c. Bakr. 1009. Bdu•ionl faidrncr fo1 • ConlXI Sa l'hnomoni: Componm1.i ih< Enlmld Ash llo<tt. Agrillll Pbnipcnnis Faimu11<. joonul of Ch<m1al Ecology JI: 104-110.

Llu. H. P.. L S. &ucr, R. Gao. T. Zhao. T. R. l'ctnc<. and R. .l Huck. 200J E.xplon10I) suoty for tmmld ash bom, Arri"' pt."i"""" (Cclrop<ttao Buprcnicbt') and its n01iu1"21 rnroiics in Cliru. Tht Grrar lAkts Enro"'°"'fur 36: 191-20-I.

I.Ju. H. L S. S.utt. D. L Milla. T. Zhao. R. C.O. L Song. Q. Lu... R. Jin. and C. G..,. 2007. S.asonal abundance of A.!fllus pJ.mrm•u (Cclrop<m: Bur=id.icl and iu rutural <11<mics 000.lll agrili (H>'ln<nopl'1'L Encyrudxl andT<112>!.chlll planil""nlli (Hu""""!"tta EulophNil<) in Chma. Biologial U>mro14!:61-71.

.\kCullough. D. G .. T. M. l'ubnd .. l C. .\nulc.ia ~nd D. Cappacn. !009• Emmlcl ..Ji bortr I.Arri'" pt."P"''" Faimuirt) (U>lcop1m: Bupr<S1idx) mraction co strcssrd or bai1td ash (fritrim.u spp.) uttS. EO\ironmmul En1ornolofl JS: 1668-16-9.

McCullough. D. G .• T. M. Pol10d, D. Cappam, and A. C. Anul"'1CL !009b. Emrrakl ash bortt (Aqi!UJ pltmpn111t1) attraction m ash lrttS ul"05<'d ~­gudhng. httbicick and •ounding. Caiudwi jounul of Fomt Roarch J9: 1))1-1341.

M.:Cullough D. G .. T. M. Pobnd. A. C. Anulc.-ia. P. l.t.i< and D. Cal'!""'.

2011. folua1ion of Arri., "'""""" conuol J>ll"iikd b. """""';" hrnzoatr .ind t\\O monic01:inoid irucaKidcs.. OM and two seasons .thtt lr<alm<m. Joonul of Economic En1omology 104: 1199-611

.lkCullough. D. G .. and R. j. M<n>dtr. !Oil SLA.11 in an uiban fo,.,,, t\~uuion of potrntial smurgics 10 Slow Ash Monality C2u.sc.J by rmmld ..Ji bom (Arri"' p/4.,pninu). ln1l'llU1ional joonul of Pest .ll=gnnrn1 SS: 9-.!l

Mcknzir K. B. Hdson, 0. Thompson. G. Otis. J. Mcfarlinr, T. Buscarini, <anJ J. \lcomng. 2010. ..\udirachun: .\n Effecti\t Sy,tcmk: lr\)(\ticKk foe Con1rol of. l.viiw ~"P"''" <Colcopitta Bup!C!tidx). jounul of E.onom• Emomolos-· 10.l: -os-T.

.lkr<>da. R. ). \ 11-. Siq;<n. \. \t. Licbhold and D. G. McCulloup.. !011. btinuting the dfmi,mm of thrtt pcxcnii.tl nwugtmtnt op1ions to skr. cht ~prtad of rmt'rJ.IJ ash bortr popuboons m l001liud outlirr mes. Carudian jounul offom1 R=rch 41: !S;-!64

\l<1uJtt. R. J .. D. G. M.Cullough and J . .\I. lkdfonl 2013. A comp•rilOI\ of girdla:l ash dr1ruion trctS and bJitnl mificu.l tr.1ps for rnmald .id-I borer Afri/111 ,U111pt1mi.s flirm1irr) dti«tion. Em·ironmmtal En1~· 42:

101--1039.

\luup•ll1. O .. X. &i. P. .\bmid•la. S. P. Rajmpu. P. Bondlo. D.A. Httllh. !010. li.hUMpt\.itk tnnscripcomioof iht nock: inusM insm poi (mmld bit bortt (Arrlw ~.,pninu). PloS On<l<IOJ: 1-12.

Pctri..r. T. R.. md R. A. HaacL 200'". Un rrmrald ash borer. AzrilUJ p1,i,,;pm,,u 1C'.okoptm: Bupra1Wd ~from logs rv.-o \Ummm afttt mfoial trco m cm? Tht GrC"Jt U.kes En10mologi~140: 92-9;.

Poland. T . .\I.: and D. G . .\lcCullowJi. !006. Erner.Id ..Ji bom: '"'"'ion oi tht urban fotot and tht thrr:u co \onh . .\merica'' ~h rtSOUM. Joom.il of Forcs1~ 10-lo 118·1!4.

Pobnd. T. \I D. G. ~kCullough. and \ . C. .\nul.-ia. !011. fa~uarion oi doohk-0.da '"!" for cm.raid ash bortt <U>kopim: Bupr"1Nil<I. jounul of Economic Emomology 1 0~: ;1-:i.531 .

Pultl>\>nn. D. l..ndT. \1. Poland. !OO'JL Hos< Scla;uoo and Fm!rngl'rtf<mkc ii .1rrl"' ~"!"'"" (Colcop1"" Bup!Clti<b<I on Ash If.,,.,., spp.J Emironmmul Emomologv 38(3): .,5.,·""6;'

Puro•>nn. D. \ and T. \I. Po~ . .!OO'Jb. The rok of olfxton· cua rn sbon­rangt mur finding by tht anmld ~ borer, Atnl1tS p/a,upnrnn Fa.irm.1.11t (Colrop1m: 8"p1cs1idac). Joum•I of illl«t lkhaHor. 220 !Oi-216.

Raimpu. I. P.. r. .ltamid.iu. D ..... Httm, P. Bondlo. and 0. \tnupall1 0. 1011. Anuok.id.tm gtnC'S or the ml(r.t!J ash borer l.Apilus p/.:mijtt11nis): scne dtmctniu1ion and nprmion profilC'. Jounul or lnsm Phn~· ,~; 81~!4

Rauschrr. I\. 2006. The !005 Mich1g.m t'mmld ~ borer ft')J"'OOfte'. an update. p. I. In I. \J.suo. R. R<.rdon. and G. Pam (U•mpiknl, Proo.mlmp of th< Emtt~ .~ llo<tt R=rch and Tcdinolog> Dc.·dopmn11 \ltturig.

P1mburgh. PA, 16-!7 'icp<. lOOi. U.S. Otpanm<111 of Agricuhure. Fom1 Sm-kc Puhl"''"" FHlTJ'.!00).16. 11oipn1 .. o. WV.

RM E. ) .. D .. l Hmiu. •nd D. R. .lmnky DR. !008. lmmpo.1f• ''"''"'n in rtsimnct to rmrrald ash bom {Colroptm; Bupres1id.tr) among ~onh American and Asian ..Ji (frmn"' 'l'P·I· EnHronrncnul En1,,,,."°s'· r 24!-!4'1.

Rodrigutt-S.OIU. C .. T. \1. Pobnd. J. R. \tiller. L. L S1tlins~. G. G. Gr.im. P. dt Cn>OI. L S..han. and I \U.Doiuld. 2006. llduriool and dmroph''">lopnl "'P"""' of ih< cmmld ..Ji t.or.:1, Agnlm pbmpcnnr" 10 inducal n:ibulcs or MAnchuri<Jn aili. Fraxmw 11Wlldil11mca. ChmlO«ology 16: -)-86.

Rodrigua-S..- C. R.. J. R. Milh T. \I. Pobnd. l \1. Kuhn. (,. \\'. °''" T. Turk. tnd ~. McKrnzir. 200~. lklu\'i<ll.irs of A<luli rmmld aJi borcr.

lrrbu "'""""" (Cokopitta 8"prnti<b<l. Th< (,m1 Uk En1omolop" 40: 1-16.

R,,JI, K. L. ~ j. Silk. P. M11v. D. Crook •. A. Khnmun. A. A. c..~. J. S""""· and T. Sarr201l .. A11r.1<1ion of .jzn/., ~"'!"'"' lc.koti1tta lluprtWcbc 10 a \'obtik Phnomone Eff((t) of Rdasr Ra1r. HOSt \'ob1ik . .ind Trip Placemtm. Environmcmal Emomolog.'" 41: 648-656.

Sill, P. ) .. 1'. R.all. D. B. Lron.. J. S.ttll<\. and). llu. !009. A conoo sn phnomonc i:umponrm oft~ anrrald .uh bom, A:nlm plampnmu Faimu1rr (U>lroptm: Bup.,1idac). Naru,.;s..,nsdulicn 96: 601.(,08.

Silk. P. ) .. K. R•all. P. \uyo, P. • .II. Laru~. G. Gr.im. D. Crool .. A. Co•<. I. F12><1 . J. D. SY.«nt'). D. B. Lyons, tl al. WI I. E\"idrn« for a \"ob.tile phcromonr m AfrilMI p/Jt1zpttt11is Fairnuirt (C.Oleoptm.: Buproiidacl that inert.ISO mf""lClion 10 •host foliar ,,,u,;k. F.n\lrorun<nW Emornolofl iO: 904-916.

Smulcy D.R.. J. J. Doccob, 0. Lu•. lOIO. Mul11plc-rca1 pro1<et10n of ash""' from tmmlJ ash bom- -.uh a singk uunk in1«1ion or mumcam bmzo.ur. and sinpMnr protea•"' .,di •• 1miibdoprid h.i"1 dimch. Aiboricul1Urt and t:iban f=s11y 36: 206-ll 1.

Sud.ling D.\t., r. c. Tobin. D. G . .lkCwlough .• nd D. A. Hmiu. !012. Combimng Txtics 10 Esiploi1 All« Effrm foi Endiation of Al"" ln><tt Popubuons. Journal of Economic En1oniology 10\: 1-13.

T ..... S.R.. and D. G . .llcCullough. 2012. Oilftt<111w pmistcll« ofblu< ... and .... 1ti1r ash folbAing trnmkbsb bortr im"aSion. Canadiln Journal of Fomt R=rch 41: 1542-1110.

Tiuatk.A. R.. D. G . .\lcCullough. andT. \I. Pobnd. !011. lnllucn«olhost""" on Clllttaid ash bo1tt l.A.!"lus p'4nrpninu fau11UJ1t} !Cclcop1m: 8"pl<Sl•bcl adul1 dtnsil)"• ~-dopmcnt, and disuibu1ion in Fritximu pmnl]fw1ir11 t~ En•iromntttul En1rnolofl 40: Ji--.~'6.

LIDA A.PHii !Lni1cd lu1cs D<p.nmcm of Agrirul1u1< Animal Pbni Hcal1h lnspmion Smkcl. !014a. USDA APHIS Pl'Q-'01• Emerald \sh Bortt Su.r1t'I· Gu.Jdi.nc. . .\\,libblt" .11: hnp:' "'-...Jphii.lb<b..~-lphru_hcalth. pl..mt_pounlO!tmm.ld_ash_b'dtJY.nkwhisuM)"-8\'iddincs.pdf (a..:co:.cJ

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LIDA APHI\ 1Ln11al Imes lkpanmm1 of .~rurt AnimB Pbn1 Hw1h ln<pmion Sm·i«). l014b. Emmld Ash Bortr, A"ibb1li~ of an fJwironmcnul As.smmrnt. A\iiliblt at: hnp;f!.,.,..,."Vl.rrgubtk:in-..gm· :~dock <11l<uil:D·APHIS-!oo--0060 (.:m><d. jun< 201•

IJSDA APHI~ (Lni1ed Si;itC'S Dqiinmcnt of Agricuhurr Animal Pfam Heilth lnspro:ion Stn·k.'t). 1010. Emerald . .\Ji Bom. Atnl11J p/Jniptnms (Flirm.iirrt B..logial Conuol Rd= and ~'"'"' Gwdtlincs. L5DA-APHl\..IR>­FS. Riwdtk. \ID. .A•·aibbk '" h11p:1 ''""•phr>.usda.go•l~ni_hw1h plani_pN_infolcm.r Bd _ .. h_ bl do• nkuds/ EAB-FlddR<l=·Guicklino pdl ,,"'-:L-mtd ~ Jufl( !0141.

llmg.X.-Y..Z.·Q. Y..g.J. R. Gould,\.-~. Zh•ng. G.·J. uu,and E.-S. Lru. 2010. Th<biolog.> mdm>logyof 1h"m.1~ash bor?T .. ~rri"'~"P"'""· inChiru )ounuJ of I""" S..1<11<c 10: 1!8. OOL i""""""'<.oig/10.128

ll'ci, X .. D. Reardon. \\. Yun. and J.-H. lun (!(!();) Em.raid a>h bor<r. Aj:ulu. plmij'lt'nni~ h:irm.Ure 1Cokopma: Buprestidk). in Chin.i: .i rnn .inJ distribution \lint\·. Acu Emomolog;ca SinK::J "-: 6 ~l)-685.

\\'ci. X.. Y llu. R. D. R<Jrdon. T.-H. Sun. \I. Lu. anJ J.-H. lun. lOO'. Biolog.> •nd rbnug.: '"'"ii cmmlJ ..Ji bom d.!"bu ,i.n.,,.niJ Faimuirtl m Ot1"' lnS«t 5'im:< 14: 36-·l'.l.

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Yu. C. 1'19!. lgrilu. nuruipoli Obcnbttj;tt. pp. 400401. h G. \im al. , Forrst lnStlt> of Clma (2.., cdmon). Qin.a Nrocrry Publi\hmg HOlbC, Beijing. Chiru.

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