Article

Co-Occurrence of the Invasive Banded and European Elm Bark Beetles (Coleoptera: Scolytidae) in North America

Jana C. Lee, Ingrid Aguayo, Ray Aslin, Gail Durham, Shakeeb M. Hamud, Bruce D. Moltzan, A. Steve Munson, José F. Negrón, Travis Peterson, Iral R. Ragenovich, Jeffrey J. Witcosky, Steven J. Seybold

Annals of the Entomological Society of America (impact factor: 1.32). 04/2009; 102(3):426-436. DOI:10.1603/008.102.0311

Source: OAI

ABSTRACT The invasive European elm bark beetle, Scolytus multistriatus (Marsham), was detected in Massachusetts a century ago, and it now occurs throughout the continental United States and southern Canada. The Asian banded elm bark beetle, Scolytus schevyrewi Semenov, was discovered in the United States in 2003, and now occurs in 28 states and the province of Alberta, Canada. Although the indigenous populations of these two species are allopatric, the invasive populations are now sympatric in North America where they co-colonize elm (Ulmus spp.) trees. A large-scale survey of these two Scolytus species was conducted with baited funnel traps, Plexiglas panel traps, and Ulmus pumila L. trap logs. Sites (four per locality) were monitored around Sacramento, CA; Reno, NV; Ogden, UT; Newcastle, WY; and Fort Collins, CO (2006-2007), and Manhattan, KS, and Columbia, MO (both only in 2007). Trap catches of S. schevrewyi relative to both Scolytus species captured from all three trapping methods at each survey site were 90 and 89% in Colorado, 90 and 83% in Wyoming, 60 and 68% in Utah, 43 and 68% in Nevada, and 11 and 13% in California (all in 2006 and 2007, respectively), and 3.3% in Kansas and 2.7% in Missouri (both only in 2007). Elevated abundances of S. schevyrewi at survey sites in Colorado and Wyoming could be the result of competitive displacement of S. multistriatus by S. schevyrewi, whose occurrence and mechanism require further study. General seasonal trends from all sites indicated peak flight in July and August for S. schevyrewi and two peaks (May-June and July-August) for S. multistriatus. Funnel traps baited with Multilure and 2-methyl-3-buten-2-ol were highly attractive to S. multistriatus, and mildly attractive to S. schevyrewi, whereas panel traps caught few beetles. The U. pumila trap logs were a more sensitive monitoring tool for detecting the presence of S. schevyrewi.

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Article: Semiochemical-Mediated Flight Strategies of Two Invasive Elm Bark Beetles: A Potential Factor in Competitive Displacement

Jana C. Lee

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ABSTRACT: A seven-state survey showed that the recently detected invasive Asian banded elm bark beetle, Scolytus schevyrewi Semenov, was abundant in areas of Colorado and Wyoming, whereas the long-established European elm bark beetle, S. multistriatus (Marsham), was not as abundant. In one of a series of studies to evaluate whether S. schevyrewi is competitively displacing S. multistriatus in their North American zone of sympatry, we characterized long-range flight responses infested or uninfested to small cut logs of American, Chinese, and Siberian elm, Ulmus americana, U. parvifolia, and U. pumila. Trials were conducted in Colorado and Wyoming to test the flight response of S. schevyrewi; in California to test the response of S. multistriatus; and in Nevada to test the responses of both species simultaneously. Studies with S. schevyrewi showed that males and females aggregated toward Ulmus spp. host volatiles but provided no evidence of a putative aggregation pheromone during the 0- to 48- or 48- to 96-h period of infestation. In contrast, S. multistriatus was attracted to U. pumila over unbaited controls, more to U. pumila infested with conspecific females than without, and more during the 48- to 96- versus 0- to 48-h period of infestation. This confirmed that male and female S. multistriatus aggregated toward host volatiles and that females produced an aggregation pheromone. In a cross-attraction study, S. schevyrewi displayed neither flight preference nor interruption to 17. pumila infested with conspecifics, heterospecifics, or a mix of both species. Response of S. multistriatus was too low to draw conclusions. Although S. multistriatus aggregates moderately to host volatiles and strongly to female-derived pheromones emitted after a few days, S. multistriatus may have a relative disadvantage by selecting elm hosts more slowly than S. schevyrewi, which aggregates very strongly to host volatiles. The differential long-range host location strategy may be one factor in a chain of behavioral events that leads to advantageous host colonization and development by S. schevyrewi.

Environmental Entomology 04/2010; · 1.56 Impact Factor
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Article: Biology of the Invasive Banded Elm Bark Beetle (Coleoptera: Scolytidae) in the Western United States

Jana C. Lee

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ABSTRACT: The banded elm bark beetle, Scolytus schevyrewi Semenov (Coleoptera: Scolytidae), native to Asia, was detected in the United States in 2003, and as of 2011 it is known to occur in 28 states and four Canadian provinces. S. schevyrewi infests the same elm (Ulmus spp.) hosts as the long-established invasive, the European elm bark beetle, Scolytus multistriatus (Marsham). Information on the basic biology of S. schevyrewi in its native range is sparse; thus, we conducted laboratory studies and field studies in Colorado and Nevada. Comparisons of flight and behavioral responses were made with co-occurring S. multistriatus. When Siberian elm, Ulmus pumila L., cut logs (bolts) were allowed to be colonized by wild populations in the field, S. schevyrewi did not differ in emergence density from 10- versus 24-cm-diameter bolts. In the laboratory, S. schevyrewi readily colonized bolts of American elm, Ulmus americana L., but not Chinese elm, Ulmus parvifolia Jacq.; Siberian peashrub, Caragana arborescens Lam.; a cherry, Prunus fontanesiana (Spach) C. K. Schneid.; or Russian olive, Elaeagnus angustifolia L. In Colorado, S. schevyrewi often landed on elm bolts between 12:00 p.m. and 4 p.m.; and near large elm trees, they were captured more frequently on sticky traps at 1.8 and 3.7 m aboveground than higher along the main stem. In Colorado/Nevada, S. schevyrewi initiated flight in April/March and ceased in October/September, whereas S. multistriatus initiated flight in April/May and ceased in October/September. In funnel trap flight assays of semiochemicals in Colorado or Nevada, S. schevyrewi had moderate responses, 3–10-fold greater than unbaited control traps, to Multilure (a commercial lure for S. multistriatus), 2-methyl-3-buten-2-ol (MB) + multistriatin, and MB + a plant extract that is included in a commercial formulation of Multilure. In contrast, S. multistriatus had a 226–259-fold greater response to Multilure than to the control. Both Scolytus species showed electroantennographic (EAG) responses to MB, racemic multistriatin, and (+)- and (-)-α-inene, with the greatest sensitivity to multistriatin. S. schevyrewi was more responsive to (-)-α-pinene than was S. multistriatus.

Annals of the Entomological Society of America 07/2011; · 1.32 Impact Factor

Data provided are for informational purposes only. Although carefully collected, accuracy cannot be guaranteed. The impact factor represents a rough estimation of the journal's impact factor and does not reflect the actual current impact factor. Publisher conditions are provided by RoMEO. Differing provisions from the publisher's actual policy or licence agreement may be applicable.

Page 1

ECOLOGY AND POPULATION BIOLOGY
Co-Occurrence of the Invasive Banded and European Elm Bark
Beetles (Coleoptera: Scolytidae) in North America
JANA C. LEE,1,2INGRID AGUAYO,3RAY ASLIN,4GAIL DURHAM,5SHAKEEB M. HAMUD,2
BRUCE D. MOLTZAN,6A. STEVE MUNSON,7JOSE´F. NEGRO´N,8TRAVIS PETERSON,9
IRAL R. RAGENOVICH,10JEFFREY J. WITCOSKY,11AND STEVEN J. SEYBOLD2
Ann. Entomol. Soc. Am. 102(3): 426Ð436 (2009)
TheinvasiveEuropeanelmbarkbeetle,Scolytusmultistriatus(Marsham),wasdetectedin
Massachusetts a century ago, and it now occurs throughout the continental United States and southern
Canada. The Asian banded elm bark beetle, Scolytus schevyrewi Semenov, was discovered in the United
States in 2003, and now occurs in 28 states and the province of Alberta, Canada. Although the indigenous
populations of these two species are allopatric, the invasive populations are now sympatric in North
Americawheretheyco-colonizeelm(Ulmusspp.)trees.Alarge-scalesurveyofthesetwoScolytusspecies
was conducted with baited funnel traps, Plexiglas panel traps, and Ulmus pumila L. trap logs. Sites (four
per locality) were monitored around Sacramento, CA; Reno, NV; Ogden, UT; Newcastle, WY; and Fort
Collins, CO (2006Ð2007), and Manhattan, KS, and Columbia, MO (both only in 2007). Trap catches of S.
schevrewyi relative to both Scolytus species captured from all three trapping methods at each survey site
were 90 and 89% in Colorado, 90 and 83% in Wyoming, 60 and 68% in Utah, 43 and 68% in Nevada, and
11and13%inCalifornia(allin2006and2007,respectively),and3.3%inKansasand2.7%inMissouri(both
onlyin2007).ElevatedabundancesofS.schevyrewiatsurveysitesinColoradoandWyomingcouldbethe
result of competitive displacement of S. multistriatus by S. schevyrewi, whose occurrence and mechanism
requirefurtherstudy.GeneralseasonaltrendsfromallsitesindicatedpeakßightinJulyandAugustfor S.
schevyrewi and two peaks (MayÐJune and JulyÐAugust) for S. multistriatus. Funnel traps baited with
Multilure and 2-methyl-3-buten-2-ol were highly attractive to S. multistriatus, and mildly attractive to S.
schevyrewi, whereas panel traps caught few beetles. The U. pumila trap logs were a more sensitive
monitoring tool for detecting the presence of S. schevyrewi.
ABSTRACT
KEY WORDS
schevyrewi
competitive displacement, ßight behavior, Scolytidae, Scolytus multistriatus, Scolytus
In the United States and Canada, ?500 exotic insect
species attack trees and shrubs, affecting forest and
urbanlandscapes(Niemela ¨ andMattson1996,Langor
et al. 2009). The European elm bark beetle, Scolytus
multistriatus Marsham [Coleoptera: Scolytidae, sensu
Wood (2007)], originates from Europe (Fig. 1a) and
wasÞrstdetectedin1909inMassachusetts(Chapman
1910). By 1970, S. multistriatus was found in most of
the contiguous United States except for Montana, Ar-
izona, and Florida (Barger and Hock 1971), but it has
since been collected in Montana in 1973 (Claßin and
Dooling1973),inArizonain1976(B.Celaya,personal
communication), and in Florida in 1997 (T. H. Atkin-
son, personal communication) (Fig. 1a). Since the
1930s,S.multistriatushasbeenknowntobeaprincipal
vector of Dutch elm disease (DED) (Readio 1935), a
disease that caused 50Ð75% mortality of pre-1930s
Americanelm,UlmusamericanaL.,populationsinthe
northernandeasternUnitedStates(BloomÞeld1979).
ThecausativeagentsofDEDarethefungiOphiostoma
himal-ulmi Brasier & M.D. Mehrota, Ophiostoma
novo-ulmi Brasier, and Ophiostoma ulmi (Buisman)
Nannf. (Harrington et al. 2001). S. multistriatus vec-
torsDEDwhensporesofthesefungiarepresentinthe
1Correspondingauthor:USDAÐARS,HorticulturalCropsResearch
Laboratory, 3420 NW Orchard Ave., Corvallis, OR 97330 (e-mail:
jana.lee@ars.usda.gov); formerly Department of Entomology, Uni-
versity of California Davis, One Shields Ave., Davis, CA 95616.
2USDAForestService,PaciÞcSouthwestResearchStation,Chem-
icalEcologyofForestInsects,720OliveDr.,SuiteD,Davis,CA95616.
3Colorado State Forest Service, 5060 Campus Delivery, Fort Col-
lins, CO 80523.
4Kansas Forest Service, 2610 Claßin Rd., Manhattan, KS 66502.
5Nevada Division of Forestry, 2478 Fairview Dr., Carson City, NV
89701.
6USDA Forest Service, Forest Health Protection, 1601 N. Kent St.,
Arlington,VA22209;formerlyMissouriDepartmentofConservation,
1110 S. College Ave., Columbia, MO 65201.
7USDA Forest Service, Forest Health Protection, 4746 South 1900
East, Ogden, UT 84403.
8USDA Forest Service, Rocky Mountain Research Station, 240
West Prospect Rd., Fort Collins, CO 80526.
9City of Newcastle, 10 West Warwick, Newcastle, WY 82701.
10USDA Forest Service, Forest Health Protection, 333 SW First
Ave., Portland, OR 97208.
11USDAForestService,ForestHealthManagement,740SimmsSt.,
Golden, CO 80401.
0013-8746/09/0426Ð0436$04.00/0 ? 2009 Entomological Society of America

Page 2

beetleÕs galleries in the phloem. New adults emerge
from pupal chambers carrying fungal spores. When
theyfeedonbranchesoflivingelms,thefungusenters
the tree and develops in the vascular tissues (Collins
et al. 1936).
The banded elm bark beetle, Scolytus schevyrewi
Semenov,originatesfromMongoliaandNorthernAsia
(references in Fig. 1a) and was Þrst detected in ßight
traps in April 2003 in Aurora, CO, and in Ogden, UT
(Negro ´n et al. 2005). Like S. multistriatus, adult S.
schevyrewi colonize elms by boring into branches and
trunks of stressed or moribund trees to feed and de-
velopintheinnerandouterbark(Wang1992).Bythe
endof2003,S.schevyrewiwasdetectedin14states,by
theendof2004insevenmorestates,andfrom2005to
2007insevenmorestates(Fig.1b).Detectionhasnot
necessarily reßected presence throughout an entire
state.Forexample,S.schevyrewiwascaughtin2003on
theeasternedgeofOregon,andasofDecember2008,
it has been caught in the northern but not the central
and southern portions of the state (Oregon Depart-
ment of Agriculture, unpublished data). Likewise, in
central and northern California it has not been re-
portedfromthecoastalurbanareasorfromthenorth-
ern Central Valley (R. L. Penrose et al., unpublished
data). The nearly simultaneous detection of S.
schevyrewiin14statesin2003stronglyimpliesthatthe
beetle likely arrived earlier but remained undetected.
This was conÞrmed when an examination of museum
collections later revealed previously collected speci-
mensofS.schevyrewi,includingtheearliestcollection
in1994fromDenver,CO(Negro ´netal.2005),andthe
ÞrstcollectionfromTexas(RandallCo.)in2002(E.G.
Riley,personalcommunication).Themodeofentryof
S. schevyrewi into North America is unknown, but
manyinvasivebarkbeetleshavebeenfoundinbarked
wood packaging materials associated with imports in-
spected at U.S. ports (Haack 2001, Lee et al. 2007).
Between 1985 and 2000, 92 interceptions of Scolytus
beetles have occurred at U.S. ports of entry, 14 spec-
imens were identiÞed to species (Haack 2006), and
none were found to be S. schevyrewi (R. A. Haack,
personal communication).
Fig. 1.
EuropeandAsia,respectively,andtheiradventivedistributionsinNorthAmerica.(b)StatesintheUnitedStatesthathavedetected
S.schevyrewisince2003andsites(dots)wherespecimenspredating2003wereidentiÞed.DistributionsbasedonMichalski(1973),
CommonwealthInstituteofEntomology(1975),WoodandBright(1992),BrightandSkidmore(1997,2002),Negro ´netal.(2005),
Lee et al. (2007), Government of Canada (2008), Langor et al. (2009), National Agricultural Pest Information System website
(http://pest.ceris.purdue.edu/pestlist.php), and personal communications from T. H. Atkinson (Florida), B. Celaya (Arizona),
J. Cena (Washington), A. Eitam (Ohio), R. Hiskes (Connecticut), D. Martin (Virginia), and E. G. Riley (Texas).
(a) Approximate native distribution of S. multistriatus (dotted delineation) and S. schevyrewi (solid delineation) in
May 2009LEE ET AL.: CO-OCCURRENCE OF INVASIVE ELM BARK BEETLES
427

Page 3

The mere presence of S. schevyrewi or any other
exotic species is not necessarily detrimental if it does
not cause harm to the economy, environment, or hu-
man health. Rather, exotic species become “invasive”
by progressing through the phases: 1) arrival; 2) es-
tablishmentÑorganism survives and reproduces; and
3) spread (Venette and Carey 1998, Liebhold and
Tobin 2008). S. schevyrewi has arrived and is estab-
lished in more than half of the land area of the con-
tinental United States and seems to have spread in
areas such as Colorado and Wyoming (Negro ´n et al.
2005, Lee et al. 2006). As invasive populations of S.
schevyrewi spread, the impacts on urban and unman-
agedlandscapesareamajorconcern.InfestationsofS.
schevyrewi alone can kill water-stressed elms, as ob-
served in China (Wang 1992) and Colorado (Negro ´n
et al. 2005). In addition, S. schevyrewi is a suspected
vector of DED because beetles collected from dis-
easedtreeswerefoundtocarrysporesofO.novo-ulmi
(Jacobi et al. 2007) although vectoring efÞciency is
still being investigated.
TheimpactandspreadofS.schevyrewiintheUnited
States will depend on its ecological interactions, par-
ticularly those with S. multistriatus, which shares the
same niche in the inner and outer bark of elms. In-
fested elm logs collected in 2003 in Fort Collins, CO,
revealed that 99% of emerging adult scolytids were S.
schevyrewi(Negro ´netal.2005),whereasS.multistria-
tus was readily found in city elm groves in the past
(Hostetler and Brewer 1976). The numerical domi-
nance of S. schevyrewi suggests that it may be com-
petitively displacing S. multistriatus, i.e., the “elimina-
tioninagivenhabitat,ofonespeciesbyanotherwhere
one possesses the identical ecological niche to the
other” (DeBach 1966). Analogously, S. multistriatus is
thought to have displaced the native elm bark beetle,
Hylurgopinus rufipes (Eichhoff), in areas where S.
multistriatus could overwinter (Lanier 1983). S. mul-
tistriatuswasconsideredtobemoreaggressiveandhas
a second generation each year, compared with the
native beetle with one generation per year. Displace-
ment of a native insect species by an exotic is unde-
sirableandrepresents33%ofthedocumentedcasesof
competitive displacement (Reitz and Trumble 2002).
If it is occurring, displacement of S. multistriatus by S.
schevyrewiwouldrepresentthedisplacementofapre-
viously established exotic by another exotic, like 55%
ofdocumentedcases.Forelmbarkbeetles,thesecond
displacement event will potentially also be undesir-
able because S. schevyrewi is causing additional dam-
agetoalreadyembattledelmpopulations,particularly
to Siberian elm, Ulmus pumila L. (Lee et al. 2007).
Unfortunately, many cases of displacement have
been accompanied by anecdotal evidence of compe-
titionwithoutempiricaldocumentation(Simberloffet
al. 1997). The current invasion of S. schevyrewi pro-
videsatimelyopportunitytobeginobtainingevidence
of displacement lacking from most post hoc studies,
particularly because this invasion may be at different
stages at different locations in the United States. Doc-
umenting the extent to which S. schevyrewi has re-
placed S. multistriatus across their respective adven-
tive ranges is a key Þrst step in exploring the
mechanism of displacement, i.e., whether displace-
ment in this case may be mediated by differential
abilities to locate hosts during ßight (J.C.L. et al.,
unpublished data), to accept hosts at close range, to
lay eggs, and the competitive ability of larvae (J.C.L.
and S.J.S., unpublished data). In this study, our ob-
jective was to quantify the relative abundance of S.
schevyrewiandS.multistriatusalongawide-scalegeo-
graphic transect that included California, Nevada,
Utah, Wyoming, Colorado, Kansas, and Missouri.
Materials and Methods
In 2006, elm bark beetles were monitored from
May/June to September at four sites each around
Sacramento, CA; Reno, NV; Ogden, UT; Newcastle,
WY; and Fort Collins, CO; in 2007, monitoring con-
tinued in the same areas with the addition of Man-
hattan, KS, and Columbia, MO (Table 1). Four sites
within each state were within a circle that was 2.4Ð98
km in diameter, and sites were selected for proximity
to standing elm trees or cut elm stem wood or
branches, and where anthropogenic disturbance of
trapswouldbelesslikely.Forbrevityintheresultsand
Þgures, we refer to the sites by their state names,
although the captures from a city or several cities of
each state do not necessarily reßect abundances
throughout the entire state. To assess populations,
threetypesoftrapswereusedsimultaneouslybecause
eachtrapmayhaveabiastowardcatchingonespecies
over another. First, a 12-unit Lindgren funnel trap
(Lindgren1983)wasbaitedwithMultilurereleasedat
0.3mg/d(PherotechInternationalInc.[nowContech
Inc., Delta, BC, Canada]), and 2-methyl-3-buten-2-ol
(Aldrich Chemical Co., Milwaukee, WI) released at
5Ð18 mg/d from 10 heat-sealed 400-?l polyethylene
Eppendorf tubes (Evergreen ScientiÞc, Los Angeles,
CA).Baitswerereplacedafter2mo.Thisbaitedßight
trap was expected to elicit a higher response from S.
multistriatus because Multilure contains the phero-
mone blend for this species. Multilure and methylbu-
tenol were also known to be mildly attractive to S.
schevyrewi (Negro ´n et al. 2005, Lee et al. 2007).
Next, a 30- by 30- by 50-cm clear Plexiglas Schmitz
panel trap (Schmitz 1984) was installed at each site
under an elm tree or next to elm debris. This trap
consisted of two Plexiglas panels slit halfway in the
center and Þtted together perpendicularly at the slits,
appearing as a Ô?Õ sign from the top view. The bottom
ofeachofthefourtrappingvaneswasconnectedtoan
inverted 3.8-liter milk jug, which was in turn con-
nected to a collection cup. This passive trap was con-
sidered to be the least biased, because it would trap
any insect ßying in the vicinity of the bole of elms or
respondingtovolatilesfromcutlogs.Funnelandpanel
traps were spaced at least 15 m apart and emptied
weekly.
ThethirdtrapwasacutSiberianelm,U.pumila,log
with ?14.1-dm2surface area and wax-sealed ends to
reduce desiccation. One trap log was laid at each site
three times each summer during May/June, July, and
428ANNALS OF THE ENTOMOLOGICAL SOCIETY OF AMERICA
Vol. 102, no. 3

Page 4

August and retrieved ?2 wk later (Table 1). Insects
were then reared from the logs at the USDA Forest
Service Chemical Ecology of Forest Insects Labora-
toryinDavis,CA,atambientconditions.Eachlogwas
placed in an individually aerated and sealed plastic
18.9-liter paint bucket (Lee et al. 2008). Emerging
adults were collected from a glass collection jar at the
bottom of each bucket every week for 2 mo because
S. schevyrewi can develop from egg to adult within
30Ð45 d (Wang 1992, Negro ´n et al. 2005), and S.
multistriatuswithin35Ð60d(Drooz1985,Cranshawet
al. 1993). After the 2-mo rearing period, logs were
stored at 4?C until they could be debarked to collect
anyremainingadults;someadultsrecolonizedthelogs
Table 1.
western United States
Description and location of trapping sites for determining the relative abundance of S. multistriatus and S. schevyrewi in the
State DescriptionLocation Latitude and longitude
Dates for ßight traps and when
trap logs were exposed in Þeld
CaliforniaPrivate residence, single U.
pumila
Shaded area, distant U.
pumila and U. americana
Green waste pile with cut
Ulmus spp.
Green waste pile, Bing
Maloney Golf Course, U.
parvifolia
Equestrian Center, U.
pumila
Green waste pile, cut
Ulmus spp.
Golf course, U. pumila
OfÞce complex, U. pumila
(2006 only)
Private residence, U.
pumila (2007 only)
Golf course, U. pumila
Campgrounds, U. americana
and U. pumila
Nature Center, U. pumila
Train depot, U. pumila
Woodland, Yolo Co. 38? 39? 53? N, 121? 46? 18? WFlight: May 26ÐOct. 4, 2006;
April 3ÐSept. 12, 2007
Trap logs: June 7Ð22, July 6Ð21,
Aug. 9Ð23, 2006; May 15Ð
June 1, July 5Ð20, July 31Ð
Aug. 15, 2007
Davis, Yolo Co.38? 32? 24? N, 121? 44? 24? W
Davis, Yolo Co.38? 32? 12? N 121? 48? 22? W
Sacramento, Sacramento Co.38? 30? 10? N, 121? 29? 41? W
Nevada Reno, Washoe Co.39? 32? 26? N 119? 48? 23? WFlight: May 24ÐOct. 3, 2006;
April 5ÐSept. 9, 2007
Trap logs: June 7Ð20, July 4Ð18,
Aug. 11Ð25, 2006; May 17Ð31,
July 6Ð20; Aug. 1Ð15, 2007
Reno, Washoe Co. 39? 31? 13? N 119? 46? 19? W
Reno, Washoe Co.
Carson City, Carson Co.
39? 30? 05? N 119? 45? 50? W
39? 07? 32? N 119? 46? 12? W
Minden, Douglas Co.38? 58? 3? N 119? 41? 31? W
UtahOgden, Weber Co.
Huntsville, Weber Co.
41? 17? 11? N, 111? 59? 15? W
41? 15? 01? N, 111? 47? 10? W
Flight: June 9ÐSept. 11, 2006;
May 18ÐSept. 11, 2007
Trap logs: June 9Ð23, July 5Ð21,
Aug. 11Ð25, 2006; May 18Ð
June 1, July 6Ð20, Aug. 3Ð17,
2007
Flight: June 15ÐSept. 13, 2006;
May 21ÐSept. 9, 2007
Trap logs: June 15Ð29, July 7Ð
20, Aug. 14Ð28, 2006; May
21ÐJune 6, July 5Ð19; Aug.
2Ð15, 2007
Ogden, Weber Co.
ClearÞeld, Davis Co.
41? 14? 42? N, 111? 59? 41? W
41? 06? 19? N, 112? 02? 19? W
Wyoming City shop, U. pumila
Private residence,
U. pumila
Private residence, U.
pumila
Green waste pile, cut
Ulmus spp.
Nursery, U. pumila (2007
only)
School, U. americana
and U. pumila (2006
only)
Research Station, U.
pumila
OfÞce complex, U. pumila
OfÞce complex, single U.
pumila
Private residence,
U. americana and
U. pumila
Private residence, U.
pumila
Private residence, U.
pumila
Private residence, U.
americana
OfÞce complex, Ulmus
spp.
Park, Ulmus spp.
OfÞce complex, Ulmus
spp.
Park, Ulmus spp.
Newcastle, Weston Co.
Newcastle, Weston Co.
43? 51? 31? N, 104? 12? 15? W
43? 51? 21? N, 104? 13? 09? W
Newcastle, Weston Co.43? 51? 09? N, 104? 11? 52? W
Newcastle, Weston Co.43? 50? 27? N, 104? 12? 59? W
Colorado Ft. Collins, Larimer Co.40? 35? 14? N, 105? 08? 42? W Flight: June 12ÐSept. 15, 2006;
May 16ÐSept. 7, 2007
Trap logs: June 13Ð28, July
10Ð26, Aug. 9Ð23, 2006; May
17ÐJune 1, July 5Ð17, Aug.
1Ð15, 2007
Ft. Collins, Larimer Co.40? 34? 23? N, 105? 04? 03? W
Ft. Collins, Larimer Co.40? 34? 06? N, 105? 05? 09? W
Lakewood, Jefferson Co.
Lakewood, Jefferson Co.
39? 43? 13? N, 105? 06? 44? W
39? 43? 28? N, 105? 07? 11? W
KansasNear Randolph, Riley Co.39? 29? 59? N, 96? 45? 22? WFlight: May 23ÐSept. 3, 2007
Trap logs: May 24ÐJune 6, July
3Ð18, Aug. 8Ð22, 2007
Near Louiseville,
Pottawatomie Co.
Manhattan, Riley Co.
39? 13? 40? N, 96? 21? 32? W
39? 09? 42? N, 96? 32? 12? W
Near Tri-County Road and
Quaker Road, Morris Co.
Columbia, Boone Co.
38? 52? 24? N, 96? 35? 51? W
Missouri38? 56? 08? N, 92? 19? 19? W Flight: May 22ÐSept. 5, 2007
Trap logs: May 22ÐJune 5,
2007, July 3Ð17; July 31ÐAug.
14, 2007
South of Columbia, Boone Co.
Columbia, Boone Co.
38? 55? 44? N, 92? 18? 40? W
38? 55? 30? N, 92? 18? 15? W
South of Columbia, Boone Co.38? 48? 45? N, 92? 15? 21? W
1Freshly cut Ulmus pumila trap logs were collected from: 1) Carson City, NV in Mar. 2006 and stored at 4?C until placement in the Þeld
in June 2006; 2) Reno, NV on June 19, 2006 for placement in the Þeld in July and Aug. 2006; and 3) Reno, NV on Mar. 8, 2007 for placement
in May, July, and Aug. 2007.
May 2009LEE ET AL.: CO-OCCURRENCE OF INVASIVE ELM BARK BEETLES
429

Page 5

after emergence and needed to be accounted for in
the assessment. Some parental beetles in good condi-
tion may have been counted as progeny because par-
ents could not be separated deÞnitively from progeny
beetles. However, this overestimation would have oc-
curred among all trap logs. The trap log served as a
measure of the expected recruitment rate of progeny
ofbothspecieswhenhostmaterialwasavailableinthe
Þeld. The trap log would likely recruit more progeny
of the earlier colonizing species if it has the compet-
itive advantage of equaling or preceding the other
species in development.
Despite their co-occurrence and the potential for
confusion, the two bark beetle species were readily
separated by external morphological characters (Ne-
gro ´netal.2005).Thesecharactersarediagnostic,even
when evaluated by the nonspecialist (LaBonte et al.
2007). For data analysis, we used the number of bee-
tles per species collected from funnel and panel traps
on a weekly basis, and the total number of beetles per
species reared from each trap log. To compare the
relative abundance of the two species, the percent-
age of S. schevyrewi adults [S. schevyrewi/(S. mul-
tistriatus?S.schevyrewi)]collectedperßighttrapor
per log was converted to a proportion and arcsine?x
transformed to normalize variances. Trap catches
withoutScolytusspp.wereexcludedfromtheanalyses.
To compare total abundance among ßight traps, the
number of S. schevyrewi or S. multistriatus captured
per trap was standardized on a daily basis (number of
beetles/sampling days between collection intervals)
and then log10(x ? 1) transformed. Among trap logs,
the total abundance of each species collected per log
was analyzed. The percentage and total S. schevyrewi
and S. multistriatus captured/reared out were ana-
lyzed with a repeated measures analysis (? ? 0.05)
with the variables state, time, and state ? time in
PROC MIXED (SAS Institute 2001). If the effect of
statewassigniÞcant,meansfromeachstateacrosstime
were compared by RyanÕs Q multiple comparison,
which has high power and controls for experiment-
wisetypeIerror(DayandQuinn1989).Becauseßight
trapping durations varied from state to state, only
samples during 21 JuneÐ13 September 2006 and 29
MayÐ7 September 2007 were incorporated in the sta-
tisticalanalysesbecauseallstatesweresimultaneously
monitored at these times. Voucher specimens from
eachlocationhavebeendepositedattheOregonState
University Arthropod Collection, Corvallis, OR (ac-
cession 00229), University of California Davis Bohart
Museum of Entomology, and the California Academy
of Sciences, San Francisco, CA.
Results and Discussion
Geographic Trends. This 2-yr monitoring survey
revealedthathigherpercentagesofS.schevyrewithan
S. multistriatus were found in sites in Colorado (CO),
Wyoming (WY), Nevada (NV), and Utah (UT) (Fig.
2). Relatively fewer S. schevyrewi than S. multistriatus
were recovered in California (CA), Kansas (KS), and
Missouri (MO) (Figs. 2 and 3). Funnel, panel, and log
trap catches were often affected by the state ? wk or
state ? mo interactions (Table 2), but general trends
overtimearediscussed.Relativeabundanceinfunnel
traps in 2006Ð2007 had the following trend: CO ?
WY ? UT ? NV ? CA ? KS ? MO (Table 2; Fig. 3a
and b). Relative abundance in panel traps had similar
trends: in 2006 CO ? WY ? UT ? NV ? CA (Fig. 3c)
and in 2007 CO ? WY ? NV ? UT ? CA ? KS ? MO
(Fig. 3d). Recruitment of beetles from trap logs
showedslightlydifferenttrendsinrelativeabundance
than the ßight traps. Nevada ranked generally higher
amongmoststatesinrecruitingahigherpercentageof
S. schevyrewi from logs: in 2006 WY ? CO ? NV ?
UT ? CA (Fig. 2e); and in 2007 CO ? NV ? UT ?
WY ? CA ? KS ? MO (Fig. 3f).
These results suggest that S. schevyrewi may be at
different stages of invasion across the western United
States with the highest abundances in Colorado and
Wyoming.Interestingly,areductioninS.multistriatus
population density seems to have occurred in Colo-
rado because S. multistriatus was recovered abun-
dantly in elm bolts in 1973 in Fort Collins, CO, and in
1974 in Denver, CO (Hostetler and Brewer 1976).
NowthehighincidenceofS.schevyrewiinFortCollins
and Lakewood (near Denver), CO, may be the result
of competitive displacement, but additional monitor-
ing over time in Colorado and other states is needed
to determine whether displacement is occurring. A
similarpatternwasfoundwiththeinvasiveeucalyptus
longhorned borers (Coleoptera: Cerambycidae) in
southern California where Phoracantha semipunctata
F., discovered in 1984 (Scriven et al. 1986), was likely
displaced by Phoracantha recurva Newman, discov-
ered in 1995 (Hanks et al. 1997). P. recurva made up
1.4% of Phoracantha spp. emerging from naturally in-
fested eucalyptus in 1996, 74% in 1997, and ?95% in
1998 (Hanks et al. 1997, Bybee et al. 2004). The ad-
vantage for P. recurva seems to lie with the suscepti-
bility of P. semipunctata to an egg parasitoid (Luhring
et al. 2004). This displacement led to establishment of
a second invasive species that was less likely to be
naturally regulated than the Þrst invasive species.
Total numbers of S. schevyrewi and S. multistriatus
collected per day in funnel and panel traps showed
similar trends as the relative abundance of the two
species, but with less separation among states. Total
capturesofS.schevyrewifollowedthistrend:infunnel
Fig. 2.
S.schevyrewiatsitesin2006Ð2007inCalifornia(CA),Nevada
(NV),Utah(UT),Wyoming(WY),andColorado(CO),and
in 2007 in Kansas (KS) and Missouri (MO).
Trapping sites (stars) and overall percentages of
430ANNALS OF THE ENTOMOLOGICAL SOCIETY OF AMERICA
Vol. 102, no. 3

Page 6

traps in 2006 WY ? CO ? UT ? NV ? CA (Fig. 4a);
infunneltrapsin2007CO?NV?UT?WY?CA?
KS?MO(Fig.4b);inpaneltrapsin2006WY?NV?
UT ? CO ? CA (Fig. 4c); and in panel traps in 2007
CO ? WY ? UT ? NV ? KS ? CA ? MO (Fig. 4d).
Total captures ßuctuated year to year; Wyoming
and Nevada varied with funnel traps, and Colorado
varied with panel traps. However, total numbers of
S. schevyrewi in funnel and panel traps were gen-
erally low, ?10 beetles per day, with a peak in ßight
during July and August (Fig. 4aÐd) similar to a 2003
survey in Lakewood, CO, where the peak captures
occurred from 15 July to 12 August 2003 (Negro ´n et
al. 2005).
TotalcapturesofS.multistriatusfollowedthistrend:
in funnel traps in 2006 NV ? CA ? UT ? CO ? WY
(Fig. 5a), in funnel traps in 2007 NV ? KS ? UT ?
CA?MO?CO?WY(Fig.5b),inpaneltrapsin2006
CA ? NV ? UT ? CO ? WY (Fig. 5c), and in panel
traps in 2007 KS ? MO ? CA ? UT ? NV ? WY ?
CO (Fig. 5d). Funnel trap captures of S. multistriatus
showed a peak in MayÐJune and a secondary peak in
JulyÐAugust, and were as high as 150/d in late July
2006 in NV (Fig. 5a and b). In 2007, panel traps in KS
caught ?50 S. multistriatus per day in early June (Fig.
5d). In Lakewood, CO, peak ßight activity of S. mul-
tistriatus was observed in mid-August (Negro ´n et al.
2005).Ina3-yrsurveyinGeorgia,S.multistriatusßight
Fig. 3.
and 2007 (b), in panel traps in 2006 (c) and 2007 (d), and recovered from trap logs in 2006 (e) and 2007 (f). Vertical dotted
linesinaÐdindicatethebeginningandendingofthesamplingperiodswhenallsiteswereanalyzedstatistically.Lettersdenote
signiÞcant differences (P ? 0.05) by RyanÕs Q on arcsine?x-transformed data; untransformed data are shown.
Mean percentage of S. schevyrewi (relative to both S. schevyrewi and S. multistriatus) in funnel traps in 2006 (a)
May 2009LEE ET AL.: CO-OCCURRENCE OF INVASIVE ELM BARK BEETLES
431

Page 7

peaked in late April/May, mid-July, and late August/
early September (Hanula and Berisford 1984).
Among trap logs, total recruitment of S. schevyrewi
from cut elm was greatest in Nevada and Colorado
during both years, and in Utah in 2007, sometimes
reaching ?1,000 within the 2-mo rearing period, or 71
beetles per dm2(Fig. 4e and f). Recruitment of S.
multistriatuswashighestinCAduringbothyears,and
in MO in 2007 (Fig. 5e and f), with ?1,000 beetles
emerging per log in these cases. Recruitment of prog-
eny for both species was always greatest among logs
exposed in the Þeld in August (Figs. 4e and f and 5e
and f). August may be the period when the highest
number of reproducing adults is present, or when
conditions are best for reproduction and develop-
ment. Interestingly, ßight of S. multistriatus toward
funnel traps peaked noticeably in late July 2006 in
Nevada (Fig. 5a) and remained steady in California
through most of the summer, whereas logs exposed in
mid-Augustinbothplacesresultedinthehighestlevel
of recruitment (Fig. 5e). This suggests that captures
from the ßight traps and trap logs may reßect activity
of different portions of a population.
Comparison of Trapping Methods. As expected,
funnel traps containing an attractant generally col-
lected more Scolytus beetles than passive panel traps.
For example, in 2006, no Scolytus spp. were collected
from eight of 261 funnel trap samples compared with
65 of 257 panel trap samples. In 2007, no Scolytus spp.
were collected from 34 of 405 funnel trap samples
compared with 108 of 405 panel trap samples. Also
expected was that funnel traps baited with the pher-
omone for S. multistriatus were more sensitive in cap-
turing S. multistriatus (as many as 150 per d) than S.
schevyrewi (no ?10 per d) (Figs. 4a and b and 5a and
b). Also, trap logs of U. pumila were much more
sensitive in detecting S. schevyrewi throughout its ad-
ventive range, even in instances when funnel and
paneltrapsfailedtodetectthebeetle.Forexample,no
S. schevyrewi were captured in funnel and panel traps
inMO;afewwerecapturedinßighttrapsinCalifornia
andKansas(Fig.4aÐd);buttraplogsrecruitedatleast
a few S. schevyrewi in Missouri, and up to 300Ð400 in
California and Kansas (Fig. 4e and f). Unlike ßight
traps, recruitment of new adults from trap logs de-
pends on the ßight and close-range responses of each
species to the log, the abilities of each species to
colonize and mate on the log surface, and the devel-
opmental capacities of their larval progeny beneath
the bark of the log. The relative sensitivity of the trap
logs as a detection tool for S. schevyrewi may be be-
cause of greater intrinsic suitability of U. pumila as a
host for S. schevyrewi than S. multistriatus. U. pumila,
likeS.schevyrewi,originatesfromcentralAsia(Moore
2003). Nevertheless, studies suggest that U. pumila is
also a highly suitable host for S. multistriatus; ?90
progenywereproducedperdm2whenS.multistriatus
wereplacedontoacutU.pumilalog(J.C.L.andS.J.S.,
unpublished data), and ?200 progeny emerged per
dm2when U. pumila logs were exposed to S. multi-
striatus populations in the Þeld (Sˇvihra and Volney
1983).Alternatively,therecruitmentbiasofU.pumila
trap logs for S. schevyrewi may have occurred because
S. schevyrewi is a better competitor in colonizing elm
hosts than S. multistriatus.
Implications. The arrival of S. schevyrewi, a second
invasiveelmbarkbeetleinNorthAmerica,represents
a rekindled threat to native and ornamental elm pop-
Table 2.
S. multistriatus captured in flight traps or emerging from trap logs
Repeated measures analysis (ANOVA) of the percentage of adult S. schevyrewi and total counts of adult S. schevyrewi and
Yr
Variable
StateWk or mo
State ? wk or
state ? mo
Trap type, variable ?transformation?
FdfPFdfPF dfP
Funnel trap
Percentage of S. schevyrewi ?arcsine?proportion?
2006
2007
2006
2007
2006
2007
20.3
21.0
3.06
5.22
26.7
6.58
4, 15
6, 21
4, 15
6, 21
4, 15
6, 21
?0.001
?0.001
0.0497
0.0020
?0.001
?0.001
1.97
1.90
2.10
3.03
2.03
5.72
12, 173
14, 246
12, 181
14, 280
12, 181
14, 280
0.0295
0.0226
0.0188
?0.001
0.0238
?0.001
1.37
1.75
1.78
1.77
1.84
3.03
48, 173
83, 246
48, 181
83, 280
48, 181
83, 280
0.0766
?0.001
0.0037
?0.001
0.0022
?0.001
Total count of S. schevyrewi ?log10(x ? 1)?
Total count of S. multistriatus ?log10(x ? 1)?
Panel trap
Percentage of S. schevyrewi ?arcsine?proportion?
2006
2007
2006
2007
2006
2007
25.9
154.8
0.95
6.63
2.08
4.66
4, 15
6, 21
4, 15
6, 21
4, 15
6, 21
?0.001
?0.001
0.463
?0.001
0.134
0.0037
1.21
0.94
1.46
3.58
1.98
3.21
12, 113
14, 173
12, 177
14, 280
12, 177
14, 280
0.282
0.518
0.145
?0.001
0.0285
?0.001
0.85
1.54
1.42
1.36
0.92
3.70
46, 113
83, 173
48, 177
83, 280
48, 177
83, 280
0.726
0.009
0.053
0.0342
0.622
?0.001
Total count of S. schevyrewi ?log10(x ? 1)?
Total count of S. multistriatus ?log10(x ? 1)?
Trap logs
Percentage of S. schevyrewi ?arcsine?proportion?
2006
2007
2006
2007
2006
2007
5.47
14.2
4.93
10.1
8.11
5.03
4, 15
6, 21
4, 15
6, 21
4, 15
6, 21
0.0064
?0.001
0.0098
?0.001
0.0011
0.0024
5.40
3.27
17.9
12.1
7.10
15.1
2, 24
2, 33
2, 30
2, 42
2, 30
2, 42
0.0116
0.0505
?0.001
?0.001
0.003
?0.001
2.07
3.07
6.41
2.52
2.62
3.97
8, 24
12, 33
8, 30
12, 42
8, 30
12, 42
0.0799
0.0053
?0.001
0.0136
0.0265
?0.001
Total count of S. schevyrewi ?log10(x ? 1)?
Total count of S. multistriatus ?log10(x ? 1)?
Only ßight trap catches from June 21 to Sept. 13, 2006 and from May 29 to Sept. 6, 2007 were included in the statistical analysis because all
states were monitored simultaneously at these times.
432ANNALS OF THE ENTOMOLOGICAL SOCIETY OF AMERICA
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Page 8

ulations in hardwood forests and urban landscapes. U.
americana has been a favored shade tree throughout
theUnitedStatesforitsfastgrowth,estheticallypleas-
ing crown, and tolerance to stress. It was distributed
in native forests throughout eastern North America,
but during the 20th Century it also became a valuable
part of the urban landscape across the United States.
The devastating DEDÐbark beetle complex killed be-
tween 50 and 75% of the pre-1930s elm population
in northeastern North America (BloomÞeld 1979;
Tainter and Baker 1996; R. J. Hauer, unpublished
data), including primarily U. americana as well as six
otherspeciesofelms(StrobelandLanier1981).How-
ever, through natural regeneration, planting of resis-
tant varieties, and good sanitation practices, U. ameri-
cana, and, to a lesser extent, U. pumila, continue to be
important urban trees, particularly in the Upper Mid-
west, the Great Plains, the Intermountain West, and
California (R. J. Hauer, unpublished data; McPherson
1998; McPherson et al. 2004; Peper et al. 2004).
Our trap logs revealed how quickly S. schevyrewi
populationscouldbuildupamongweakenedelmtrees
or debris. In Colorado, trap logs with ?14.1-dm2sur-
face area recruited up to 1,000 S. schevyrewi adults
within 2 mo (Fig. 4e and f). The Intermountain West
and Midwest may be vulnerable because U. pumila is
one of the few shade trees present in communities
suchasLovelock,Winnemucca,andBattleMountain,
NV;Edgemont,SD;andLusk(personalobservations),
andNewcastle,WY,where333infestedelmtreeswere
Fig. 4.
and recovered from trap logs in 2006 (e) and 2007 (f). Vertical dotted lines in aÐd indicate the beginning and ending of the
sampling periods when all sites were analyzed statistically. Letters denote signiÞcant differences (P ? 0.05) by RyanÕs Q on
log10(x ? 1)-transformed data; untransformed data are shown.
Mean number of S. schevyrewi in funnel traps in 2006 (a) and 2007 (b), in panel traps in 2006 (c) and 2007 (d),
May 2009LEE ET AL.: CO-OCCURRENCE OF INVASIVE ELM BARK BEETLES
433

Page 9

removedin2004(Leeetal.2006).InFortCollins,CO,
U. americana and U. pumila make up 18.5 and 9.5% of
canopy cover, respectively (McPherson et al. 2004).
Loss of these trees could lead to a substantial increase
in energy use and reduction of air quality. Minnesota
hasreported9,706,10,869,16,617,and36,537elmtrees
removed because of DED from 2001 to 2004, respec-
tively, corresponding to a 12% increase from 2001 to
2002, 53% increase from 2001 to 2003, and 220% in-
crease from 2001 to 2004 (Burks 2005). Although the
cause for this increase is not known, the role of S.
schevyrewi may be of interest. In laboratory and Þeld
cage trials, S. schevyrewi reared from diseased elms
were allowed to feed on elms and found to transmit
the pathogen to the new wound (Koski and Jacobi
2007). This suggests that S. schevyrewi may transfer
DED, although the vectoring efÞciency is still being
investigated. However, if S. schevyrewi is a less efÞ-
cient vector of DED than S. multistriatus, which has
yettobedetermined,adisplacementeventmayeven-
tually be beneÞcial for elm populations in the United
States.
Our 2-yr monitoring survey indicates how predom-
inant S. schevyrewi is in parts of Colorado and Wyo-
ming, with substantial populations in Utah and Ne-
vada, and low populations thus far in California,
Kansas, and Missouri. Although the point or points
where S. schevyrewi was introduced to the United
States are unknown, the data collected here and pre-
vious collection data are consistent with S. schevyrewi
Fig. 5.
and recovered from trap logs in 2006 (e) and 2007 (f). Vertical dotted lines in aÐd indicate the beginning and ending of the
sampling periods when all sites were analyzed statistically. Letters denote signiÞcant differences (P ? 0.05) by RyanÕs Q on
log10(x ? 1)-transformed data; untransformed data are shown.
Mean number of S. multistriatus in funnel traps in 2006 (a) and 2007 (b), in panel traps in 2006 (c) and 2007 (d),
434ANNALS OF THE ENTOMOLOGICAL SOCIETY OF AMERICA
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Page 10

establishingfounderpopulationsinColoradoandWy-
oming. The earliest pinned specimens were from Col-
orado and New Mexico, which supports the hypoth-
esis of founderpopulations
surrounding areas. We might better understand the
invasion pattern of S. schevyrewi by conducting pop-
ulation genetic studies to elucidate how closely re-
lated the populations are over the adventive geo-
graphic range, and whether one or more points of
introduction occurred.
inColoradoand
Acknowledgments
WethankT.H.Atkinson(DowAgroSciences,Austin,TX;
retired), B. Celaya (OfÞce of State Forester, Phoenix, AZ),
J. Cena (Washington State Department of Agriculture,
Olympia,WA),A.Eitam(USDAÐAPHIS,Pickerington,OH),
R. J. Hauer (University of Wisconsin, Stevens Point, WI), R.
Haack (USDA Forest Service [FS], East Lansing, MI), R.
Hiskes (Connecticut Agricultural Experiment Station, New
Haven, CT), J. R. LaBonte (Oregon Department of Agricul-
ture, Salem, OR), D. Martin (Virginia Department of Agri-
culture&ConsumerServices,Richmond,VA),R.L.Penrose
(California Department of Agriculture, Sacramento, CA),
and E. G. Riley (Texas A & M University, College Station,
TX) for unpublished data on the distribution of S. multistria-
tusandS.schevyrewiandportinterceptionrecords.Wethank
the following people for assistance: [California] P. Jiros ˇ
(USDA FS, Davis); T. Eißer, E. Espiritu, S. Genito, S. Hishi-
numa, L. Shell, T. Sze-To, C. Wang, T. Young (University of
California, Davis); S. Howe (Bing Maloney Golf Course,
Sacramento); K. Daane (University of California, Berkeley);
[CO] S. Costello, D. Dean, J. Popp (USDA FS); Johnson
Investments; Colorado State Experiment Station, Harris El-
ementary School, Ft. Collins; P. McCusker (Colorado FS);
[KS]K.Bomberger,R.Hauck,T.Rhodes,S.Zimmer(Kansas
FS);Z.andP.Bomberger(Dwight);[MO]D.Brandt(Mon-
tana Department. Conservation); J. and M. Stevens (Colum-
bia); [NV] M. Ghiggeri (Nevada Division of Forestry); S.
Churchillo (Reno Parks & Recreation); V. Markussen (Car-
sonCity);T.Jennings(Brookside);D.Kennedy(UNRValley
Road);J.Romwall(MercuryAirCenter);R.Inman,M.Reitz
(Reno); [UT] L. Dunning, B. Bentz (USDA FS); M. Mc-
Kinley (Ogden Nature Center); Freeport Center; J. Tracy
(Ben Lomond); C. Sibbernsen (Unita-Wasatch-Cache NF
Ogden RD); [WY] Cheyenne East High School; L. Koch
(Wyoming State Forestry Division); and B. Terry, C. Allison
(Newcastle). Funding was provided by USDA NRI CSREES
grant 2006-35302-16611, USDA Forest Service Special Tech-
nology Development Program grant R2-2004-05, PaciÞc
Northwest Region R6 Forest Health Protection, and the
PaciÞc Southwest and Rocky Mountain Research Stations.
We also thank D. Ullman, S. Padgett, and L. Beneze (UC
Davis, Department of Entomology) for administrative sup-
port.
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Received 9 October 2008; accepted 23 January 2009.
436 ANNALS OF THE ENTOMOLOGICAL SOCIETY OF AMERICA
Vol. 102, no. 3

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Keywords

28 states

continental United States

Elevated abundances

Fort Collins

Funnel traps baited

indigenous populations

invasive populations

large-scale survey

mildly attractive

panel traps

Plexiglas panel traps

S. multistriatus

S. schevyrewi

Scolytus multistriatus

Scolytus schevyrewi Semenov

southern Canada

three trapping methods

two Scolytus species

Ulmus spp

United States

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