Trapping Phyllophaga spp. (Coleoptera: Scarabaeidae: Melolonthinae) in the United States and Canada using sex attractants

Abstract

The sex pheromone of the scarab beetle, Phyllophaga anxia, is a blend of the methyl esters of two amino acids, L-valine and L-isoleucine. A field trapping study was conducted, deploying different blends of the two compounds at 59 locations in the United States and Canada. More than 57,000 males of 61 Phyllophaga species (Coleoptera: Scarabaeidae: Melolonthinae) were captured and identified. Three major findings included: (1) widespread use of the two compounds [of the 147 Phyllophaga (sensu stricto) species found in the United States and Canada, males of nearly 40% were captured]; (2) in most species intraspecific male response to the pheromone blends was stable between years and over geography; and (3) an unusual pheromone polymorphism was described from P. anxia. Populations at some locations were captured with L-valine methyl ester alone, whereas populations at other locations were captured with L-isoleucine methyl ester alone. At additional locations, the L-valine methyl ester-responding populations and the L-isoleucine methyl ester-responding populations were both present, producing a bimodal capture curve. In southeastern Massachusetts and in Rhode Island, in the United States, P. anxia males were captured with blends of L-valine methyl ester and L-isoleucine methyl ester.

Full text

Trapping Phyllophaga spp. (Coleoptera:
Scarabaeidae: Melolonthinae) in the United States
and Canada using sex attractants.
Paul S. Robbins1,*, Steven R. Alm2, Charles. D. Armstrong3, Anne L.
Averill4, Thomas C. Baker5, Robert J. Bauernfiend6, Frederick P.
Baxendale7, S. Kris Braman8, Rick L. Brandenburg9, Daniel B. Cash10,
Gary J. Couch11, Richard S. Cowles12, Robert L. Crocker13, Zandra D.
DeLamar14, Timothy G. Dittl15, Sheila M. Fitzpatrick16, Kathy L.
Flanders14, Tom Forgatsch17, Timothy J. Gibb18, Bruce D. Gill19, Daniel O.
Gilrein20, Clyde S. Gorsuch21, Abner M. Hammond22, Patricia D.
Hastings23, David W.Held24, Paul R. Heller5, Rose T. Hiskes12, James L.
Holliman25, William G. Hudson26, Michael G. Klein27, Vera L. Krischik28,
David J. Lee29, Charles E. Linn, Jr.1, Nancy J. Luce4, Kenna E.
MacKenzie30, Catherine M. Mannion31, Sridhar Polavarapu32,†, Daniel A.
Potter33, Wendell L. Roelofs1, Brian M. Royals9, Glenn A. Salsbury34,
Nathan M. Schiff35, David J. Shetlar36, Margaret Skinner37, Beverly L.
Sparks38, Jessica A. Sutschek39, Timothy P. Sutschek39, Stanley R.
Swier40, Martha M. Sylvia41, Neil J. Vickers42, Patricia J. Vittum4,
Richard Weidman23, Donald C. Weber43, R. Chris Williamson44 and
Michael G Villani1,†
1Cornell Univ., New York State Agric. Experiment Station, Geneva, NY psr1@cornell.edu,
cel1@cornell.edu, wlr1@cornell.edu
2Univ. of Rhode Island, Kingston, RI stevealm@uri.edu
3Univ. of Maine, Orono, ME charlesa@umext.maine.edu
4Univ. of Massachusetts, Amherst MA aaverill@ent.umass.edu, nluce@psis.umass.edu,
pvittum@ent.umass.edu
5Pennsylvania State Univ., University Park PA tcb10@psu.edu, prh@psu.edu
6Kansas State Univ., Manhattan, KS rbauernf@oz.oznet.ksu.edu
7Univ. of Nebraska, Lincoln NE fbaxendale1@unl.edu
8Georgia Experiment Station, Griffin, GA kbraman@griffin.uga.edu
9North Carolina State Univ., Raleigh, NC rick_brandenburg@ncsu.edu, brian_royals@ncsu.edu
10 Franklinville, NY dbcash@cnyti.com
11 Cornell Univ. Cooperative Extension, Middletown, NY gjc15@cornell.edu
12 Connecticut Agricultural Experiment Station, Windsor, CT richard.cowles@po.state.ct.us,
13 Texas Department of Agriculture, Austin, TX robert.crocker@agr.state.tx.us
14 Auburn Univ., Auburn, AL zd289coupe@yahoo.com, flandkl@auburn.edu
15 Ocean Spray Cranberries, Babcock, WI tdittl@oceanspray.com
16 Agriculture & Agri-Food Canada, Agassiz, British Columbia, Canada fitzpatricks@agr.gc.ca
17 Bandon, OR osonegro@peoplepc.com
18 Purdue Univ., West Lafayette, IN gibb@purdue.edu
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Journal of Insect Science: Vol. 6 | Article 39 1

19 Center for Plant Quarantine Pests, Ottawa, Canada gillbd@inspection.gc.ca
20 Cornell Univ. Cooperative Extension, Riverhead, NY dog1@cornell.edu
21 Clemson Univ., Clemson, SC csgorsuch@att.net
22 Louisiana State Univ., Baton Rouge, LA ahammond@agcenter.lsu.edu
23 Rutgers Univ. Cooperative Extension, New Brunswick, NJ hastings@aesop.rutgers.edu,
weidman@rcre.rutgers.edu
24 Mississippi State Univ, Biloxi, MS dwh56@ra.msstate.edu
25 Alabama Agricultural Experiment Station, Marion Junction, AL jhollima@acesag.auburn.edu
26 Univ. of Georgia, Tifton, GA wghudson@uga.edu
27 Ohio State Univ., Wooster, OH klein.10@osu.edu
28 Univ. of Minnesota, St. Paul, MN krisc001@tc.umn.edu
29 New York State Tree Nursery, Saratoga Springs, NY djlee@gw.dec.state.ny.us
30 Agriculture & Agri-Food Canada, Kentville, Nova Scotia, Canada mackenziek@agr.gc.ca
31 Univ. of Florida, Homestead, FL cmannion@mail.ifas.ufl.edu
32 Rutgers Univ., Blueberry and Cranberry Research Center, Chatsworth, NJ
33 Univ. of Kentucky, Lexington, KY dapotter@uky.edu
34 Kansas Department of Agriculture, Greensburg, KS gsalsbury@sbcglobal.net
35 USDA Forest Service, Stoneville, MS nschiff@asrr.arsusda.gov
36 Ohio State Univ., Columbus, OH shetlar.1@osu.edu
37 Univ. of Vermont, Burlington, VT margaret.skinner@uvm.edu
38 Univ. of Georgia, Athens, GA bsparks@uga.edu
39 Tarpon Springs, FL jessie_sutschek@yahoo.com
40 Univ. of New Hampshire, Durham, NH stan.swier@unh.edu
41 Univ. of Massachusetts Cranberry Experiment Station, Wareham,MN martys@umext.umass.edu
42 Univ. of Utah, Salt Lake City, UT vickers@biology.utah.edu
43 USDA-ARS, Beltsville, MD weberd@ba.ars.usda.gov
44 Univ. of Wisconsin, Madison, WI rcwillie@entomology.wisc.edu
†Deceased - Sridhar Polavarapu and Michael Villani are greatly missed by family, friends, and
colleagues.
Abstract
The sex pheromone of the scarab beetle, Phyllophaga anxia, is a blend of the methyl esters of two
amino acids, L-valine and L-isoleucine. A field trapping study was conducted, deploying different
blends of the two compounds at 59 locations in the United States and Canada. More than 57,000 males
of 61 Phyllophaga species (Coleoptera: Scarabaeidae: Melolonthinae) were captured and identified.
Three major findings included: (1) widespread use of the two compounds [of the 147 Phyllophaga
(sensu stricto) species found in the United States and Canada, males of nearly 40% were captured]; (2)
in most species intraspecific male response to the pheromone blends was stable between years and over
geography; and (3) an unusual pheromone polymorphism was described from P. anxia. Populations at
some locations were captured with L-valine methyl ester alone, whereas populations at other locations
were captured with L-isoleucine methyl ester alone. At additional locations, the L-valine methyl
ester-responding populations and the L-isoleucine methyl ester-responding populations were both
present, producing a bimodal capture curve. In southeastern Massachusetts and in Rhode Island, in the
United States, P. anxia males were captured with blends of L-valine methyl ester and L-isoleucine
methyl ester.
Resumen
La feromona sexual del escarabajo, Phyllophaga anxia, es una mezcla de los ésteres metílicos de dos
aminoácidos, L-valina y L-isoleucina. Se condujo un estudio de campo usando diferentes mezclas de los
dos componentes en 59 sitios de Estados Unidos y Canada. Más de 57,000 machos de 61 especies de
Phyllophaga fueron capturados e identificados. Tres de los resultados más importantes incluyen: (1) el
extenso uso de los dos componentes [de las 147 especies de Phyllophaga (sensu stricto), en Estados
Unidos y Canada, fueron capturados machos de cerca del 40% de ellas.]; (2) para la mayoría de las
especies, la respuesta intraespecífica de los machos a las combinaciones de los dos aminoácidos fue
consistente entre años diferentes, y en todos los sitios geográficos; y (3) un inusual polymorfismo de la
feromona fue descrito para P. anxia. Poblaciones de algunos sitios fueron atrapados sólo con valina,
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mientras que poblaciones de otros sitios fueron atrapados sólo con isoleucina. También se encontraron
sitios donde las poblaciones responden a ambos componentes, valina e isoleucina, produciendo una
curva de captura bimodal. En el sureste del estado de Massachusetts y en Rhode Island, en Estados
Unidos, machos de P. anxia fueron atrapados en trampas con mezclas de valina e isoleucina.
Correspondence: *psr1@cornell.edu
Received 12.10.2006 | Accepted 14.2.2006 | Published: 15.11.2006
Copyright: Creative Commons Attribution 2.5
ISSN: 1536-2442 | Volume 6, Number 39
Cite this paper as:
Robbins PS, Alm SR, Armstrong CD, Averill AL, Baker TC, Bauernfiend RJ, Baxendale FP, Braman SK,
Brandenburg RL, Cash DB, Couch GJ, Cowles RS, Crocker RL, DeLamar ZD, Dittl TG, Fitzpatrick SM, Flanders KL,
Forgatsch T, Gibb TJ, Gill BD, Gilrein DO, Gorsuch CS, Hammond AM, Hastings PD, Held DW, Heller PR, Hiskes
RT, Holliman JL, Hudson WG, Klein MG, Krischik VL, Lee DJ, Linn Jr. CE, Luce NJ, MacKenzie KE, Mannion CM,
Polavarapu S, Potter DA, Roelofs WL, Royals BM, Salsbury GA, Schiff NM, Shetlar DJ, Skinner M, Sparks BL,
Sutschek JA, Sutschek TP, Swier SR, Sylvia MM, Vickers NJ, Vittum PJ, Weidman RB, Weber DC, Williamson RC,
Villani MG. 2006. Trapping Phyllophaga spp. (Coleoptera: Scarabaeidae: Melolonthinae) with sex attractants in
the United States and Canada. 124pp. Journal of Insect Science 6:39, available online: insectscience.org/6.39.
Introduction
The scarab beetle genus Phyllophaga (sensu lato)
is one of the largest genera of animals in the
United States (Woodruff and Beck 1989),
encompassing 203 described species in 8
subgenera, including 147 species (and 8
subspecies) in the subgenus Phyllophaga (sensu
stricto), 39 species in Listrochelus, 7 species in
Phytalus, 3 species in Cnemarachis (all
non-native and confined to south Florida), 3
species in Eugastra, 2 species in Tostegoptera, 1
species in Chlaenobia, and 1 species in Triodonyx
(Evans 2003, Smith and Evans 2005). Their
striking genitalic morphology, first described in
the late 19th century (Smith 1888), continues to
be the most important taxonomic character used
to separate species in this group (Luginbill and
Painter 1953). See Woodruff and Beck (1989) and
Woodruff and Sanderson (2004) to view excellent
scanning electron microscopy images of
Phyllophaga genitalia.
The economic importance of this genus relates
principally to the root feeding habits of the larvae,
commonly called white grubs. Larvae of various
species of Phyllophaga have been recorded
feeding on crops that include, but are not limited
to, nursery stock (Andre 1937), corn (Bessin
1999), commercial turfgrass (Brandenburg and
Villani 1995; Vittum et al. 1999), cranberries
(Dunn and Averill 1996; Eck 1990; Franklin
1950), sugarcane (Gordon and Anderson 1981),
sweet potato (Hammond et al. 1997), and pasture
(Luginbill and Painter 1953). True to the Greek
origin of their generic name (Phyllo-leaf +
phaga-eat), adult Phyllophaga in very large
flights have been known to defoliate stands of
trees. Although adults of some Phyllophaga
species are apparently host specific, most are
polyphagous (Luginbill and Painter 1953).
P. anxia (LeConte) is the most widely distributed
Phyllophaga species in North America (Luginbill
and Painter 1953; Woodruff and Beck 1989). Two
genitalic morphs are described in this species, the
northern form and the southern form (Luginbill
and Painter 1953; Woodruff and Beck 1989). The
first sex pheromone described from the genus
Phyllophaga was identified from virgin P. anxia
adults dug in mid-April from a cranberry bog in
Carver, Massachusetts, before their May flight.
The female-produced sex pheromone was
determined to be a 75/25 blend of L-valine methyl
ester and L-isoleucine methyl ester (Zhang et al.
1997). L-isoleucine methyl ester was first
elucidated from Holotrichia parallela (Leal et al.
1992), an Asian melolonthine species. Holotrichia
is regarded by some as being inseparable
taxonomically from the Nearctic Phyllophaga
(Saylor 1937; Saylor 1939).
L-valine methyl ester and L-isoleucine methyl
ester are unusual pheromone compounds in that
their precursors are the essential amino acids,
L-valine and L-isoleucine. These essential amino
acids are available only via sequestration from
food plants fed on by the larvae, or perhaps from
endo-symbionts. Our future investigations will
hopefully determine the source of these amino
acids. For the most recent overview of beetle
semiochemicals see Francke and Dettner (2005).
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Since P. anxia is a common species throughout
the Northeast, the pheromone was deployed in
the field near Geneva, New York, in 1996. P. anxia
males were captured with this blend, as expected.
However, another species of Phyllophaga,P.
futilis, was also captured in the traps in much
smaller numbers. Our interest was piqued by the
P. futilis catches because sex pheromones are
generally regarded as species-specific mate
recognition signals, although studies have
demonstrated varying degrees of pheromone
specificity between closely related species
(Roelofs 1978), as well as pheromone
polymorphism in geographically separated
conspecifics such as Agrotis segetum (Wu et al.
1999), Ostrinia nubilalis (Glover et al. 1991; Klun
and Cooperators 1975), and Hemileuca
eglanterina (McElfresh and Millar 2001). Since P.
anxia is a common species throughout most of
North America, this finding presented an
opportunity to examine the response specificity of
different populations of P. anxia, as well as
responses of other Phyllophaga species, over a
large geographic region.
Materials and Methods
Vane traps baited with various blends of L-valine
methyl ester and L-isoleucine methyl ester were
deployed at 59 different locations in the U.S. and
Canada during the years 1996–2001 (Figure 1). At
each of these locations, traps were maintained for
one to four seasons. Tables 1a and 1b list the trap
locations, years during which traps were
deployed, and a brief note about the habitat. The
trapping sites in Carver, Lakeville, and Plympton,
Massachusetts; Chatsworth, New Jersey;
Babcock, Wisconsin; Lincolnville Center, Maine;
Aggasiz, British Columbia; and Bandon, Oregon,
were chosen because the traps could be located
adjacent to cranberry acreage. Researchers
involved in studies of Phyllophaga infesting turf,
pasture, nursery, or other commodities
maintained some trapping sites. Other sites were
selected because it was likely that they might
harbor different Phyllophaga species from those
in other geographic areas and where the lures had
never been tested.
When blends are referred to in this study, it is in
the ratio of L-valine methyl ester/L-isoleucine
methyl ester. In 1996, five blends were deployed,
including 100% L-valine methyl ester, 65/35,
50/50, 35/65, and 100 % L-isoleucine methyl
ester. In 1997 and 1998, 95/5 and 5/95 blends
were added to the array. In 1999, 2000, and 2001,
the eight blends tested included 100% L-valine
methyl ester, 90/10, 80/20, 60/40, 40/60,
20/80, 10/90, and 100% L-isoleucine methyl
ester. In 1996, the lures were produced in our own
laboratory by loading 5 mm rubber stopper septa
(Thomas Scientific, www.thomassci.com/
index.jsp) with 3 mg each of various blends using
hexane as the solvent. From 1997 to 2001, Dr.
A.C. Oehlschlager of ChemTica Internacional S.A.
(San Jose, Costa Rica, www.chemtica.com)
generously supplied the rubber septa lures for the
tests. During that time, the lures were loaded with
4 mg of the various blends. At each location a
control trap with a blank septum also was
deployed.
In 1996 and 1997, the traps used in the study were
either Trécé Japanese beetle vane traps (Trécé
Incorporated, www.trece.com/) or Fuji Flavor
Company vane traps (Fuji Flavor Company,
www.fjf.co.jp/). Beginning in the 1998 season,
vane traps were fabricated in the laboratory from
three-liter soda bottles and 4 mm white
corrugated plastic (Figure 2). When removed
from the field during the winter, these traps lasted
up to three field seasons.
Traps were set in the field 15–20 meters apart and
at heights of 1–2 meters. The traps were checked
and re-randomized one to three times each week.
Captured beetles were bagged and frozen, or
infrequently preserved in ethanol. Plastic bags or
bottles marked with the catch date and blend
were shipped at the completion of the trapping
period to Geneva, NY, for identification.
Phyllophaga species identifications were assigned
using a number of published sources (Luginbill
and Painter 1953;Ratcliffe 1991;Riley 1988, Saylor
1939; Saylor 1940; Woodruff and Beck 1989),
comparisons with Phyllophaga species in the
Cornell University insect collection, and
consultations with and verifications by E. Richard
Hoebeke (Cornell University, Ithaca, NY), Dr.
Paul Lago (University of Mississippi, University,
MS), Edward C. Riley (Texas A & M University,
College Station, TX), and William B. Warner
(Farnam Companies, Inc., Phoenix, AZ). In 1998,
Dr. Robert Crocker (then at Texas A & M
University, Dallas, TX) did identifications of the
Texas catch and sent the results to Geneva NY. In
2000 and 2001, Dr. Robert Bauernfiend (Kansas
State University, Manhattan, KS) did the same for
the Manhattan, Kansas catches. Whenever
possible, a series of each species from the various
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locations was pinned for later vouchering in the
Cornell University insect collection.
Results and Discussion
General observations
The following outline condenses the large number
of figures and tables found in this publication into
general groupings for the convenience of the
reader.
Table or Figure Description of Table or Figure
Tables 1a and 1b Trapping locations, years, and description of trapping sites
Tables 2a and 2b Trap catches, arranged alphabetically by species
Tables 3a and 3b Trap catches by the number caught of each species,
descending
Tables 4a and 4b Trap catches by number of sites where each species was
caught
Tables 5-97 Catches at individual locations
Table 98 Synchronically flying species captured by similar sex
attractant blends
Figure 1 Geographic locations of all trapping sites
Figure 2 Photo of trap
Figures 3-63 Maps of species distributions and where caught in this study
Figures 64-126 Male sex attractant capture curves, arranged alphabetically
by species
Figure 127 Phyllophaga male sex attractant response curves, general
observations
Figure 128 Sympatric and synchronic flights to different sex attractant
blends
Figure 129 Sympatric and asynchronic flights to the same sex attractant
blend
Figure 130 Sympatric and synchronic flights to the same sex attractant
blend
Figure 131 Seasonal flights of all species - Kansas, Manhattan, 2001
Figure 132 Seasonal flights of all species - Kentucky, 2000
Figure 133 Seasonal flights of all species - Massachusetts, Amherst, 1999
Figure 134 Male P. anxia capture curve - L-valine methyl ester
responders
Figure 135 Male P. anxia capture curve - L-isoleucine methyl ester
responders
Figure 136 Male P. anxia capture curve - bimodal site
Figure 137 Male P. anxia capture curve - blend responders
Figure 138 Map of all sites where P. anxia was found and male capture
curves
A total of 57,129 Phyllophaga individuals were
examined and identified in the course of this
study. The 215 captured females represented only
0.38% of the total catch, suggesting that the
attractants do not function as aggregation
pheromones in any of the species. A total of 145
males were counted from the control traps,
amounting to only 0.25% of the total catch.
No Phyllophaga beetles were captured in Aggasiz,
British Columbia; Bandon, Oregon; or
Homestead, Florida. In British Columbia and
Oregon, the traps were located at cranberry bogs.
A light trap operated at the Oregon cranberry bog
site also captured no Phyllophaga. Despite
extensive sampling for root weevils in northwest
commercial cranberry bogs, no white grubs have
been reported (D. C. Weber, unpublished data).
Moreover, when the Phyllophaga species
distribution maps in Luginbill and Painter (1953)
are examined, only four species of Phyllophaga
are found listed from Oregon. Regarding the
Homestead, Florida site, Woodruff (1961)
indicates that very few of the Florida Phyllophaga
species occur in Miami or in the Florida Keys. In
the same publication, he points out that “The soil
in the Miami area is extremely shallow and
underlain by öolitic limestone and, in general, is
not a good soil for white grubs.” Only 4 of the 42
species of Phyllophaga recorded from Florida
have been collected in that area.
Tables 2a and 2b list alphabetically the
Phyllophaga species captured, the number of
males captured in each species, the number of
discrete locations at which they were captured,
and the total number of site-years for each species
[site-years = (site A x number of years that species
was captured at that site) + (site B x number of
years that species was captured at that site) +
.......]. Tables 3a and 3b are sorted by descending
catch numbers, beginning with the Phyllophaga
species caught in the greatest number. Tables 4a
and 4b are sorted by site, listing in descending
order the number of sites at which a particular
species was recorded.
Of all site-years when and where beetles were
captured, only two (NJ Chatsworth #2, 1999
Table 59 and UT Salt Lake City, 1999, Table 91)
recorded a single species of Phyllophaga during a
flight season. The average number of species
captured during a site-year was 4.15. Three sites
(AL Auburn, 1998, Table 6; KS Manhattan #1,
2000, Table 27; and KS Manhattan #2, 2001,
Table 28) recorded more than 10 species of
Phyllophaga captured during the flight season.
The species that was taken in the greatest number
(20,480), in the greatest number of sites (33), and
in the greatest number of site-years (54) was P.
anxia. This is not surprising given that, as was
indicated previously, it is the most widespread
species of Phyllophaga in North America.
Luginbill and Painter (1953) list it as occurring in
every state in the United States except Arizona,
California, Florida, Nevada, West Virginia, and
Wyoming. They also report P. anxia from all ten
Canadian provinces. Woodruff and Beck (1989)
have subsequently reported it from Florida. W. B.
Warner (personal communication) has examined
a specimen of P. anxia from California and has
reports of P. anxia in Arizona.
Males of ten other Phyllophaga species were also
captured in numbers >1000. These include P.
futilis,P. congrua,P. crassissima,P.
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praetermissa,P. gracilis,P. hirtiventris,P.
ephilida,P. rubiginosa,P. rugosa, and P. fusca.
Males of 13 species were captured in numbers
>100 but <1000 and males of 15 species were
captured in numbers >10 but <100 (Tables 3a and
3b). Eighteen species of Phyllophaga were
recorded at ≥5 locations and 8 species were
recorded at ≥11 locations (Tables 4a and 4b).
Geographical distributions and range
extensions
Geographical distribution maps of Phyllophaga
species captured during this study are found in
Figures 3–63, arranged alphabetically by species.
Shaded areas in these figures indicate the
geographical ranges conveyed by Luginbill and
Painter (1953) or Woodruff and Beck (1989) [P.
(Phytalus)georgiana and P. (Phytalus)obsoleta
only].
Most captures of Phyllophaga species recorded
during this study were found within the
geographical species distributions reported by
Luginbill and Painter (1953). There are, however,
several range extensions to report. The following
species were found in locations in addition to
those reported by Luginbill and Painter: P.
curialis (Figure 14), P. drakei (Figure 17), P.
forbesi (Figure 20), P. foxii (Figure 22), P. futilis
(Figure 25), P. gracilis (Figure 29), P. gracilis var.
angulata (Figure 30), P. hirtiventris (Figure 33),
P. longispina (Figure 40), P. lota (Figure 41), P.
marginalis (Figure 43), P. (fraterna)
mississippiensis (Figure 46), P. postrema (Figure
50), P. praetermissa (Figure 51), P. quercus
(Figure 53), and P. taxodii (Figure 59). Riley
(1988) had previously noted range extensions of
P. forbesi,P. quercus, and P. taxodii into
Louisiana.
Changes in population levels from year to
year within sites
Numbers of beetles of a particular species
changed dramatically from year to year at the
same trapping location. For example, traps were
maintained at the Auburn, Alabama site during
the 1997, 1998, and 1999 seasons for
approximately the same time period each year.
The numbers of P. gracilis captured declined
from 1123 in 1997, to 710 in 1998, and to 98 in
1999 (Tables 5, 6, 7). In Lexington, Kentucky,
from 1999 to 2000, the numbers of each of the six
species captured more than doubled (Tables 29,
30).
Factors affecting the size of the population
include not only weather during the flight period,
but soil moisture and tilth conditions suitable for
oviposition, egg hatch, and growth of larvae
during the previous year or years that it takes for
development to adults. Quality and quantity of
larval host plants, soil textural characteristics, and
species’ preferences for soil types also factor into
population size and distributions (Katovich et al.
1998). Changes in population size of other species
at other trapping sites may be seen in Tables
5–97. Species captured at a particular site
changed from year to year as well, indicating that
multiyear studies will yield a more realistic
picture of population sizes and species
distributions than will a single year of captures.
Male captures with sex attractants
The most important finding of the present study
was the demonstration of the extensive use of the
methyl esters of L-valine and L-isoleucine as sex
attractants in the mate recognition systems of the
Phyllophaga. In 56 discrete locations across the
US and Canada, during 94 observation periods
(some locations were trapped for multiple years),
61 species of Phyllophaga were captured. The
overwhelming majority (58) of these species are
found in the Phyllophaga (sensu stricto)
subgenus. Since there are 147 species in this
subgenus in America north of Mexico (Evans
2003; Smith and Evans 2005), 39% of the species
in this group were captured in traps during the
course of this study.
Male trap captures are graphically illustrated in
Figures 64–126. The figures are arranged
alphabetically by Phyllophaga species. Within
each species figure, graphs are arranged
alphabetically by state or province abbreviation.
The graphs demonstrate three general patterns of
species-specific male responses to a particular
blend or group of blends. The three general
patterns are displayed in Figure 127. First, some
species, such as P. vehemens, flew primarily to the
100/0 L-valine methyl ester/L-isoleucine methyl
ester lure and were sensitive to increasing
amounts of L-isoleucine methyl ester in the other
blends, its presence significantly reducing
captures1.P. congrua, however, had a broader
response profile and was captured not only with
the 100 % L-valine methyl ester lure, but also with
blends containing 10%–20% L-isoleucine methyl
ester (Figure 71). A similar situation was seen in
responses of male P. hirtiventris (Figure 96). The
second case (Figure 127) involved species such as
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P. forbesi that were captured primarily with the
100% L-isoleucine methyl ester lure. Increasing
titers of L-valine methyl ester resulted in reduced
or no captures2. In the third case (Figure 127),
some species of Phyllophaga, such as P.
glabricula, required the presence of both
compounds before captures occurred3. An
examination of these male- response curves
reveals that, whereas certain species had a rather
broad response range to the L-valine methyl
ester/L-isoleucine methyl ester blends,4others
exhibited response curves occupying a narrower
range of blends5.
Specificity of response over time and space
A striking aspect of the intra-specific male flight
responses to the sex attractants was their
consistency between years and across geographic
locations. Several species were recorded at only
one site, but were captured at that site for two
consecutive years6. The response profiles for each
species in both years at those sites were nearly
identical.
The majority of Phyllophaga species were
recorded at more than one site (Tables 4a and
4b). As with the across-years comparison above,
the intra-specific male-response curves from
different geographic locations are similar as well.
For instance, only five specimens of P. balia were
captured during this study, but they were
captured at three different locations (Figure 6)
and all with the 100% L-isoleucine methyl ester
lure (Figure 68). Nearly 7000 P. congrua were
captured at eight different sites (Figure 9) and all
sites exhibited similar response curves (Figure
71). Comparable results are seen when other
species are examined7.
Some species were captured at only one location
and only during a single year, but multiple catches
over time in the same or nearby blends furnish a
series of independent observations that support
responses to a particular lure despite the small
numbers. For instance, six specimens of P. davisi
Langston were captured in Marion Junction,
Alabama in 1999, all in traps baited with the 100%
L-isoleucine methyl ester lure (Figure 77) on six
different dates between 4/5 and 4/28. P. davisi is
a species that Luginbill and Painter (1953)
indicate is rare, with “Only a few specimens seen.”
In Auburn, AL in 1998 two specimens of P.
diffinis (Blanchard) were captured with the 100%
L-valine methyl ester lure (Figure 78) - one taken
on 4/16 and one taken on 4/21.
Another uncommon species, P. (Phytalus)
georgiana Horn, was captured in Tifton, Georgia,
in 2000. Two individuals of this species flew to
the 100% L-valine methyl ester lure (Figure 89),
one on 7/17 and one on 7/21. Woodruff and Beck
(1989) report that no adult host plants are
recorded, the larva is undescribed, and the life
cycle is unknown. This species is one of seven
North American species in the subgenus Phytalus.
Members of this subgenus can be discriminated
from the Phyllophaga (sensu stricto) by their cleft
tarsal claws.
P. mariana is reported as a very rare species
(Luginbill and Painter 1953). Four specimens
were captured in Tifton, Georgia, in 2000 and
2001, on four different dates, in traps baited with
the 90/10 or 80/20 L-valine methyl
ester/L-isoleucine methyl ester blends (Figure
107).
Two specimens of P. taxodii Langston were
captured in Louisiana in 1997 in the 35/65
L-valine methyl ester/L-isoleucine methyl ester
blend (Figure 122), one on 7/3 and the other on
8/15. Luginbill and Painter (1953) list this species
as uncommon, having been captured only in AL
and MS. However, Riley (1988) reports that this
species is not frequently taken at lights and that
fair numbers have been captured in
flight-intercept traps 50 feet above the ground in
cypress stands. Riley concludes that a lack of light
trap catches gives the impression of rarity but that
the method of collection may be more important.
Similarly, R. J. Bauernfiend in Manhattan, Kansas
indicated that in many years of light trapping he
had never captured P. sylvatica, and was
surprised to capture 104 individuals of this
species during two years of sex attractant trapping
(Figure 121).
Of some interest is the capture of both P.gracilis
and P.gracilis variety angulata in traps baited
with 100% L-isoleucine methyl ester (Figures 92
and 93). Woodruff and Beck (1989) indicate that
“The exact status of this form (angulata) awaits
further study”. The different forms of the male
genitalia easily separate these populations of P.
gracilis. Photos of the two genitalic forms can be
seen in Luginbill and Painter (1953). The two
forms are sympatric over a large range (Figures
29 and 30). Langston (1927) provides illustrations
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of the genitalia indicating what appears to be a
form intermediate between P. gracilis and P.
gracilis variety angulata.
Intra-location species interactions
The location tables (Tables 5–97) list the blends
presented at each site in a particular trapping year
and the numbers of Phyllophaga captured with
each blend. At each site, the listed species are, by
definition, sympatric. At some sites, sympatric
species displayed asynchronous flight patterns,
including species captured days, weeks, or in
some cases, months apart. At other sites, different
species of Phyllophaga were synchronic as well as
sympatric, but may or may not have been
captured with the same sex attractant blends. The
distinction of whether males of different species
were or were not captured in the same blends is
important because it can aid in identifying
locations where inter-specific competition for
pheromonal space may be occurring. Congeneric
males flying to the same sex attractant blends
indicate situations where there is potential for
inter-specific mating interactions that afford
opportunities for investigation into reproductive
isolation involving additional species-specific sex
attractant compounds, close-range mating
behavior, and/or mating at different times.
Each of the three scenarios (1. sympatric and
synchronically flying species captured using
different blends; 2. sympatric and asynchronically
flying species captured using the same blend; 3.
sympatric and synchronically flying species
captured using the same blend) were encountered
in the course of this study, individually as well as
in combination at different study sites. The
following examples illustrate the three scenarios.
- In Lexington, Kentucky, in 1999, both P. futilis
and P. rugosa flew synchronically between
5/10 and 7/7, but were captured in traps
baited with different blends of L-valine
methyl ester/L-isoleucine methyl ester
(Figure 128).
- In Lincoln, Nebraska, in 1999, both P. vehemens
and P. crenulata were captured in the trap
baited with the 100% L-valine methyl ester,
but their flight periods were separated by 18
days during which no males of either species
were captured. P. vehemens flew from 5/16
through 5/25, whereas P. crenulata flew from
6/12 through 7/5 (Figure 129).
- In Amherst, Massachusetts, in 1999, both P.
anxia and P. longispina flew to the 100%
L-isoleucine methyl ester lure during the
period between 5/18 and 6/15 (Figure 130).
This latter scenario is the most engaging
because it is in this case that the potential for
conflict in terms of pheromonal space arises.
From the male capture data at the various
trapping sites, many examples of different species
flying synchronically, sympatrically, and to the
same blends or blend groupings have been
documented. However, nothing is known about
whether these species fly at different times of the
night or how close-range courtship behaviors are
involved in mate recognition. Figures 131–133
display detailed information relating male
response specificity curves and time of flight from
three sites that demonstrate the complexity of
interactions that can occur over a season. Figure
131 (KS Manhattan #2, 2001), demonstrates that
P. inversa and P. vehemens flew synchronically to
the 100% L-valine methyl ester lure. Similarly,
there is potential for interaction between P.
rubiginosa and P. sylvatica (middle L-valine
methyl ester/L-isoleucine methyl ester blends), P.
crassissima and P. fusca (lower L-valine methyl
ester/L-isoleucine methyl ester blends), and P.
futilis and P. bipartitia (100% L-isoleucine
methyl ester lure). In the July and August flights
at the same site, P. glabricula,P. affabilis, and P.
ephilida were captured synchronically, but with
different blends. Figure 132 (KY Lexington, 2000)
shows that P. futilis and P. hirticula flew
synchronically to the 100% L-isoleucine methyl
ester lure, whereas P. ephilida was captured with
the same lure, but later in the season. In Figure
133 (MA Amherst, 1999), synchronic species P.
forsteri,P. fraterna, and P. fusca displayed
overlapping male response curves to L-valine
methyl ester/L-isoleucine methyl ester mixes,
whereas P. anxia,P. longispina, and P. drakei
also flew synchronically to the 100% L-isoleucine
methyl ester lure.
Table 98 outlines, in a briefer format, other
trapping locations and the species involved where
inter-specific attraction might occur as a result of
competition for pheromonal space. However, the
possibility of interactions postulated by the
overlap of the male response curves in time and
space may not necessarily reflect the reality in the
field in that females may or may not have a
narrower range of sex pheromone blend
production than is suggested by what the males
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are capable of responding to. The potential for
inter-specific interactions might be predicted with
greater accuracy by combining analysis of blend
ratios produced by a number of individual females
with knowledge of male captures by various
blends and determining if male response curves
overlap congeneric female production curves.
Interspecific copulation between Phyllophaga
species has been reported in the literature. Fattig
(1944) reports (p. 26) that he collected a male P.
hirticula copulating with a female P. anxia.
Referencing the data from the present study, P.
hirticula males were captured in traps baited with
the 100% L-isoleucine methyl ester lure (Figure
95). P. anxia males of both the northern and the
southern genitalic form were also captured in
traps baited with the 100% L-isoleucine methyl
ester lure (Figures 66a, 66b, 66c, 66d, 66e and
67). It is likely that the male P. hirticula flew
upwind following an L-isoleucine methyl ester
plume to find not a conspecific female P.
hirticula, but a congeneric female P. anxia. P.
hirticula and P. anxia males possess genitalia
whose cuticular structures are exceedingly
different (see images in Woodruff and Beck,
1989). Their soft sac structures are very different
as well (P.S. Robbins, personal observation).
Their vestitures also differ, P. anxia being
glabrous and P. hirticula being hirsute. Although
interspecific genitalic differences could and
sometimes do play a role in reproductive isolation
of some taxa (Eberhard 1985, Sota and Kubota
1998), genitalic differences clearly did not prevent
copulation from taking place in this case.
Copulation does not inevitably lead to fertilization
or production of offspring (Eberhard 1996), and
similarly, attraction to a congeneric Phyllophaga
female does not inevitably lead to copulation. In a
study from Costa Rica, Eberhard (1993), reporting
on the copulatory behavior of several species of
Phyllophaga, states of P. vicina, that “Early in the
evening solitary females rested immobile and
apparently emitted an attractant, as males arrived
in flight from downwind. The pheromone
apparently also attracted males of P. valeriana, as
on three occasions I saw one or more males of this
species hover near a female P. vicina (beetles
were captured to verify their species identity).
One P. valeriana male landed on the female, then
immediately took flight and left, suggesting that a
second cue, possibly on the beetle’s surface, was
used to distinguish species identity. Contact
pheromones may be used in the melolonthine
genus Macrodactylus (Eberhard 1992).” These
observations indicate that more studies are
needed to clarify the role of morphology and/or
contact pheromones in close-range mate
recognition in Phyllophaga. Eberhard (1993)
reports that “secondary sexual modifications of
the sculpturing of the front legs, the ventral
bristles, and the overall leg length of male
Macrodactylus may function as courtship devices
prior to and during copulation”. Although the
Phyllophaga are renowned for their extravagant
and often asymmetric genitalic morphology, male
hind tibial spurs also often assume unique
configurations that provide excellent taxonomic
characters for species assignments (Luginbill and
Painter 1953; Woodruff and Beck 1989; Woodruff
and Sanderson 2004). The role tibial spurs play in
close-range mate recognition or copulatory
courtship in the Phyllophaga is unclear. Males of
some Phyllophaga species also have extensive
ventral bristles or possess unique morphological
characters on the venter, similar to
Macrodactylus (P.S. Robbins, personal
observation).
Intraspecific variation in male response: P.
anxia
The consistency of intra-specific male responses
over space and time that was discussed earlier
contrasts with the extensive variation noted in the
male-response curves of P. anxia to the various
blends of L-valine methyl ester/L-isoleucine
methyl ester sex attractants (Figures 66a, 66b,
66c, 66d, and 66e). The variations demonstrated
in the male-response profiles of P. anxia are of
four general forms:
- Those profiles from locations where the males
flew primarily to L-valine methyl ester alone
(Figure 134 and Figures 66a, 66b, 66c, 66d,
and 66e including CT Vernon, 1999; NY
Bellona, 1999 and 2000; NY Saratoga
Springs, 1999 and 2000; NY Warwick, 1999
and 2000; NY Waterloo #1, 1998 NY
Waterloo #1, 1999 and 2000; NY Waterloo
#2, 1998; ON Woodlawn, 1999 and 2000; PA
State College, 1999 and 2000, WI Babcock,
1999 and 2000)
- Those profiles from locations where the males
primarily flew to L-isoleucine methyl ester
alone (Figure 135 and Figures 66a, 66b, 66c,
66d, and 66e including MA Amherst, 1999
and 2000; NH Madbury, 1999 and 2000).
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- Those profiles from locations where males flew
to L-valine methyl ester alone or L-isoleucine
methyl ester alone on the same night in the
same place (Figure 136 and Figures 66a, 66b,
66c, 66d, and 66e, ME Lincolnville Center,
1999 and 2000; NY Franklinville, 1999; VT
Burlington, 1999 and 2000).
- Those profiles from locations where the males
were mainly captured in traps baited with
blends of L-valine methyl ester and
L-isoleucine methyl ester (Figure 137 and
Figures 66a, 66b, 66c, 66d, and 66e, MA
Carver, 1996, 1998, 1999, and 2000; MA
Lakeville, 1999 and 2000; MA Plympton #1,
1996; MA Plympton #2, 1996; RI Kingston,
1999 and 2000).
All the P. anxia males from Figures 134 to 137, as
well as all other P. anxia males in Figures 66a,
66b, 66c, 66d, and 66e are of the northern
genitalic form (Luginbill and Painter 1953;
Woodruff and Beck 1989). Male P. anxia of the
southern genitalic form (Luginbill and Painter
1953; Woodruff and Beck 1989) were captured
exclusively with L-isoleucine methyl ester alone
(Figure 67). They were captured in both a smaller
number of locations (8 vs.25) and in much
smaller numbers (20 vs. 20,640) (Table 2a) than
P. anxia males of the northern genitalic form.
Unpublished data from studies conducted at the
Franklinville, NY, site suggest that L-isoleucine
methyl ester responding P. anxia males (both
northern and southern genitalic forms) are more
sensitive to the presence of L-valine methyl ester
than are L-valine methyl ester responding males
to the presence of L-isoleucine methyl ester. Male
captures in the L-isoleucine methyl ester-baited
traps may have been suppressed by the presence
of the L-valine methyl ester at the trap sites.
The manner in which the different forms of the P.
anxia male sex pheromone response profiles are
distributed across North America reveals
important information. The five locations yielding
response profiles (10 site-years) from P. anxia
males that were captured in blends of both
L-valine methyl ester and L-isoleucine methyl
ester are found only in southeast Massachusetts
and Rhode Island (see Figures 66a, 66b, 66c, 66d,
and 66e including all the MA Carver, MA
Lakeville, MA Plympton #1, MA Plympton #2 and
RI Kingston sites). Surrounding these five
locations are trapping sites to the west (as far west
as Wisconsin) and to the north (as far north as the
provinces of Nova Scotia and Ontario) that
represent those populations of P. anxia males that
responded to only L-valine methyl ester or to only
L-isoleucine methyl ester, but did not require a
blend of the two for capture (Figure 138). The
male response curves generated by the beetle
captures at those trapping sites yield a
distribution map that is patchy in terms of
unequal distributions of the two populations.
Some sites harbor only one of the two
populations, while other sites hold both, thus
generating a bimodal distribution curve.
A detailed examination (including the soft sacs) of
male individuals of the L-valine methyl ester
responding populations, the L-isoleucine methyl
ester responding populations, and the blend
responding populations of P. anxia revealed no
character that could be used to differentiate the
populations. Further work is planned that will use
DNA sequence data from both mitochondrial and
nuclear genes to generate gene genealogies with
which to document genetic relationships within
and among the three races across the US.
Microsatellite markers will also be used to
characterize allele frequencies in natural
populations and consequently estimate the extent
of gene exchange among different pheromone
races where they occur together and between
geographically isolated populations of single
pheromone races.
Intra-specific variation in male response:
P. fraterna
A second instance of intra-specific variation in
male response to blends presented was noted at
the State College, Pennsylvania, trapping site in
1999 and 2000. Male individuals of a species
determined as P. fraterna were captured by two
blend groupings in both years (Tables 80 and 81
and Figures 85 a, 85b, and 86). In 1999, 18 males
were captured with the 0/100 L-valine methyl
ester/L-isoleucine methyl ester blend, whereas 13
males were captured with the L-valine methyl
ester/L-isoleucine methyl ester mixtures. In
2000, 19 males were captured with the 0/100
L-valine methyl ester/L-isoleucine methyl ester
blend whereas 5 males were captured with the
L-valine methyl ester/L-isoleucine methyl ester
mixtures. Phyllophaga fraterna from locations
other than Pennsylvania were captured
exclusively in the blends of L-valine methyl
ester/L-isoleucine methyl ester (Figures 85a and
85b).
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Summary and Conclusions
This study demonstrates the extensive use of the
methyl esters of L-valine and L-isoleucine in the
mate recognition systems of a widely distributed
and speciose taxon. Since each trapping site is a
snapshot of activity over a very restricted area, it
is compelling that nearly 40% of the Phyllophaga
(sensu stricto) species in America north of Mexico
were captured, despite all the possible locations
that were not trapped.
Consistency in male response among
geographically separated populations of
conspecifics was demonstrated in numerous
examples. This was useful to document in itself
because it provided information concerning sex
attractant use in a taxon that had never before
been investigated in this fashion. However, this
information also served, perhaps more
importantly, to contrast and highlight the unusual
geographic variation in male response to sex
attractants between various populations of P.
anxia and in a more minor way, P. fraterna.
This study also documents the interspecific
interactions between Phyllophaga species,
through both literature citations and capture
results. These interactions may be as benign as
attraction to congeneric females (Eberhard 1993),
with presumably little or no accompanying loss of
fitness, or may involve copulation and perhaps
exchange of genetic material (Fattig 1944),
resulting in a more expensive or even fatal
conclusion. Interactions between species are of
great interest because of their connection to both
species concepts and hybridization studies
(Arnold 1997, Claridge et al. 1997, Rand and
Harrison 1989).
Since this trapping study was terminated in 2001,
additional research on sex pheromones of the
Phyllophaga has been accomplished that extends
the understanding of mate finding in this large
genus. This research will have bearing on the
phylogenetic relationships of this group as well.
The sex pheromone of Phyllophaga crinita was
identified as methyl 2-(methylthio)benzoate
(Robbins et al. 2003). Interestingly, Coca-Abia
(2002) has recently resurrected the genus
Trichesthes and removed P. crinita as well as a
number of other species from the Phyllophaga
(sensu stricto) to Trichesthes. Among the species
moved to Trichesthes were P. lenis and P. tristis,
two species that have also been captured in large
numbers using methyl 2-(methylthio)benzoate
(P.S. Robbins, unpublished data).
The sex pheromone of Phyllophaga
(Tostegoptera)lanceolata was identified as
L-leucine methyl ester (Nojima et al. 2003).
Tostegoptera is a small sub-genus of the
Phyllophaga consisting of only two species,
Phyllophaga (Tostegoptera)lanceolata and
Phyllophaga (Tostegoptera)squamipilosa. Both
species have been captured using L-leucine
methyl ester and field tests have demonstrated
that L-valine and L-isoleucine methyl esters
function as antagonists to Phyllophaga
(Tostegoptera)lanceolata males.
Notes
1see Figure 99, P. inversa; Figure 102, P.
latifrons; Figure 106, P. marginalis; Figure 126,
P. vehemens
2see Figure 68, P. balia; Figure 69, P. bipartita;
Figure 74, P. crenulata; Figures 79a and 79b, P.
drakei; Figure 82, P. forbesi; Figures 88a, 88b,
88c, and 88d, P. futilis; Figure 92, P. gracilis;
Figure 94, P. hirsuta; Figure 95, P. hirticula;
Figure 103, P. longispina; Figure 113, P.
postrema; Figure 124, P. ulkei
3see Figure 64, P. aemula; Figure 65, P. affabilis;
Figure 72, P. corrosa; Figures 73a and 73b, P.
crassissima; Figures 83a, 83b, and 83c, P.
forsteri; Figures 85a and 85b, P. fraterna; Figures
87a, 87b, 87c, and 87d, P. fusca; Figure 90, P.
glaberrima; Figure 91, P. glabricula; Figure 104,
P. lota; Figure 107, P. mariana; Figure 108, P.
micans; Figure 112, P. perlonga; Figure 114, P.
praetermissa; Figure 116, P. quercus; Figure 117,
P. rubiginosa; Figures 118a and 118b, P. rugosa;
Figure 121, P. sylvatica; Figure 123, P. torta;
Figure 125, P. uniformis
4see Figures 73a and 73b, P. crassissima; Figures
87a, 87b, 87c, and 87d, P. fusca; Figure 117, P.
rubiginosa
5see Figure 64, P. aemula; Figure 65, P. affabilis;
Figures 83a, 83b, and 83c, P. forsteri; Figures 85a
and 85b, P. fraterna; Figures 118a and 118b, P.
rugosa; Figure 125, P. uniformis
6see Figure 64, P. aemula; Figure 65, P. affabilis;
Figure 69, P. bipartita; Figure 72, P. corrosa;
Figure 91, P. glabricula; Figure 107, P. mariana;
and Figure 125, P. uniformis
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7see Figures 11, 73a and 73b, P. crassissima;
Figures 12 and 74, P. crenulata; Figures 17, 79a
and 79b, P. drakei; Figures 20 and 82, P. forbesi;
Figures 21, 83a, 83b, and 83c, P. forsteri; Figures
23, 85a and 85b, P. fraterna; Figures 24, 87a,
87b, 87c, and 87d, P. fusca; Figures 25, 88a, 88b,
88c, and 88d, P. futilis; Figures 27 and 90, P.
glaberrima; Figures 29 and 92, P. gracilis;
Figures 31 and 94, P. hirsuta; Figures 32 and 95,
P. hirticula; Figures 33 and 96, P. hirtiventris;
Figures 36 and 99, P. inversa; Figures 40 and
103, P. longispina; Figures 41 and 104, P. lota;
Figures 43 and 106, P. marginalis; Figures 45 and
108, P. micans; Figures 47 and 110, P. nitida;
Figures 49 and 112, P. perlonga; Figures 50 and
113, P. postrema; Figures 51 and 114, P.
praetermissa; Figures 53 and 116, P. quercus;
Figures 54 and 117, P. rubiginosa; Figures 55,
118a and 118b, P. rugosa; Figures 58 and 121, P.
sylvatica; Figures 60 and 123, P. torta; Figures 61
and 124, P. ulkei; and Figures 63 and 126, P.
vehemens
Acknowledgements
This study would not have been accomplished
without the participation of my co-authors. I am
deeply grateful for their persistence and attention
to detail. Donna Boyce, Joe Ogrodnick, and Rob
Way of Communication Services here at the New
York State Agricultural Experiment Station in
Geneva provided the artistic guidance and
technical input to produce the excellent figures
seen in this publication. Thanks go to Nancy
Consolie, my lab partner in the Villani soil insect
lab at Cornell University for more than 13 years.
Her research suggestions and invariable good
humor added to this project. Marta Luisa del
Campo, Anuar Morales, and Rebecca Smyth of
Cornell University and Miguel Ángel Morón of the
Instituto de Ecologia in Xalapa, Mexico, all
contributed to making the Spanish abstract clear
and understandable. My dissertation committee
at Cornell University, consisting of Drs. Cole
Gilbert, Rick Harrison, Charlie Linn, Wendell
Roelofs, and Mike Villani, were patient and
encouraging during this long study. A special note
of thanks to Mike Villani who did not live to see
this work published. Thanks, Mike, for
encouraging me to follow the muses.
References
Andre F. 1937. White grub devastations in Iowa nurseries.
Journal of Economic Entomology 30: 615-618.
Arnold ML. 1997. Natural Hybridization and Evolution.
Oxford University Press.
Bessin RT. 1999. White grub control in no-till field corn, 1998.
In Saxena KN, editor. Arthropod Management Tests. 207.
Lanham, Maryland: Entomological Society of America.
Brandenburg RL, Villani MG, editors. 1995. Handbook of
Turfgrass Insect Pests. Lanham, Maryland:
Entomological Society of America.
Claridge MF, Dawah HA, Wilson MR, editors. 1997. Species:
The Units of Diversity. London: Chapman and Hall.
Coca-Abia MM. 2002. Reestablishment of the genus
Trichesthes Erichson, 1847 (Coleoptera: Scarabaeidae,
Melolonthinae) based on phylogeny. Journal of the New
York Entomological Society 110: 95-114.
Dunn JB, Averill AL. 1996. Survey of soil scarab insects in
Massachusetts. Cranberries 60: 9-12.
22–23.
Eberhard WG. 1985. Sexual Selection and Animal Genitalia.
Harvard University Press.
Eberhard WG. 1992. Species isolation, genital mechanics, and
the evolution of species-specific genitalia in three species
of Macrodactylus beetles (Coleoptera, Scarabaeidae,
Melolonthinae). Evolution 46: 1774-1783.
Eberhard WG. 1993. Copulatory courtship and morphology of
genitalic coupling in seven Phyllophaga species
(Coleoptera: Melolonthidae). Journal of Natural History
27: 683-717.
Eberhard WG. 1993. Functional significance of some
secondary sexual characters in three species of
Macrodactylus (Coleoptera: Melolonthidae).
Coleopterists Bulletin 47: 53-60.
Eberhard WG. 1996. Female control: sexual selection by
cryptic female choice. Princeton University Press.
Eck P. 1990. The American Cranberry. Rutgers University
Press.
Evans AV. 2003. A checklist of the New World chafers
(Coleoptera: Scarabaeidae: Melolonthinae). Zootaxa 211:
1-458.
Fattig PW. 1944. The Phyllophaga or May beetles of Georgia.
Atlanta, GA: Emory University.
Francke W, Dettner K. 2005. Chemical signalling in beetles. In
Schulz S, editor. Topics In Current Chemistry. 85–166.
Berlin: Springer-Verlag.
Franklin HJ. 1950. Cranberry insects of Massachusetts.
Bulletin 445. Parts II-VII. Massachusetts Agricultural
Experiment Station.
Journal of Insect Science | www.insectscience.org ISSN: 1536-2442
Journal of Insect Science: Vol. 6 | Article 39 12

Glover TJ, Knodel JJ, Robbins PS, Eckenrode CJ, Roelofs WL.
1991. Gene flow among three races of European corn
borers (Lepidoptera: Pyralidae) in New York state.
Environmental Entomology 20: 1356-1362.
Gordon RD, Anderson DM. 1981. The species of Scarabaeidae
(Coleoptera) associated with sugarcane in south Florida.
Florida Entomologist 64: 119-138.
Hammond A, Story RN, Murray MJ, McCown CR, Ring D.
1997. Evaluation of selected soil and foliar insecticides on
sweet potatoes for control of white grubs and banded
cucumber beetles, 1995. In Saxena KN, editor. Arthropod
Management Tests. 169–170. Lanham, Maryland:
Entomological Society of America.
Katovich K, Levine SJ, Young DK. 1998. Characterization and
usefulness of soil-habitat preferences in identification of
Phyllophaga (Coleoptera: Scarabaeidae) larvae. Annals of
the Entomological Society of America 91: 288-297.
Klun JA, Cooperators . 1975. Insect sex pheromones:
Intraspecific pheromonal variability of Ostrinia nubilalis
in North America and Europe. Environmental
Entomology 4: 891-894.
Langston J. 1927. Phyllophaga of Mississippi. Technical
Bulletin No. 15. Mississippi Agricultural Experiment
Station.
Leal WS, Matsuyama S, Kuwahara Y, Wakamura S, Hasegawa
M. 1992. An amino acid derivative as the sex pheromone
of a scarab beetle. Naturwissenschaften 79: 184-185.
Luginbill P, Painter HR. 1953. May Beetles of the United States
and Canada. Technical Bulletin No. 1060. United States
Department of Agriculture.
McElfresh JS, Millar JG. 2001. Geographic variation in the
pheromone system of the saturniid moth Hemileuca
eglanterina.Ecology 82: 3505-3518.
Nojima S, Robbins PS, Salsbury GA, Morris BD, Roelofs WL,
Villani MG. 2003. L-leucine methyl ester: The
female-produced sex pheromone of the scarab beetle
Phyllophaga lanceolata.Journal of Chemical Ecology 29:
2439-2446.
Rand DM, Harrison RG. 1989. Ecological genetics of a mosaic
hybrid zone: Mitochondrial, nuclear, and reproductive
diferentiation of crickets by soil type. Evolution 43:
432-449.
Ratcliffe BC. 1991. The Scarab Beetles of Nebraska. Bulletin of
the University of Nebraska State Museum.
Riley EG. 1988. The Phyllophaga of Louisiana (Coleoptera:
Scarabaeidae). Department of Entomology, M.S. Thesis.
Louisiana State University.
Robbins PS, Crocker RL, Nojima S, Morris BD, Roelofs WL,
Villani MG. 2003. Methyl 2-(methylthio)benzoate: the
unique sulfur-containing sex pheromone of Phyllophaga
crinita.Naturwissenschaften 90: 517-520.
Roelofs WL. 1978. Threshold hypothesis for pheromone
perception. Journal of Chemical Ecology 4: 685-699.
Saylor L. 1937. Necessary changes in status of important
Rhizotrogid genera (Col. Scarabaeidae). Revista de
Entomologia 7: 318-322.
Saylor LW. 1939. Revision of the beetles of the Melolonthine
subgenus Phytalus of the United States. Proceedings of
the United States National Museum 86: 157-167.
Saylor LW. 1940. Revision of the scarabaeid beetles of the
Phyllophagan subgenus Listrochelus of the United States,
with discussion of related subgenera. Proceedings of the
United States National Museum 89: 59-130.
Smith ABT, Evans AV. 2005. A supplement to the checklist of
the New World chafers (Coleoptera: Scarabaeidae:
Melolonthinae) with notes on their tribal classification.
Zootaxa 1032: 29-60.
Smith JB. 1888. Notes on some Lachnosterna of temperate
North America, with descriptions of new species.
Proceedings of the United States National Museum 11:
481-525.
Sota T, Kubota K. 1998. Genital lock-and-key as a selective
agent against hybridization. Evolution 52: 1507-1513.
Vittum PJ, Villani MG, Tashiro H. 1999. Turfgrass Insects of
the United States and Canada. 2nd Edition. Cornell
University Press.
Woodruff RE. 1961. A Cuban May beetle, Phyllophaga
(Cnemarachis)bruneri, in Miami, Florida (Coleoptera:
Scarabaeidae). Gainesville: Florida Department of
Agriculture and Consumer Services. Division of Plant
Industry.
Woodruff RE, Beck BE. 1989. The scarab beetles of Florida
(Coleoptera: Scarabaeidae) Part II. The May or June
beetles (genus Phyllophaga). Arthropods of Florida and
Neighboring Land Areas. 13. Gainesville: Florida
Department of Agriculture and Consumer Services.
Division of Plant Industry.
Woodruff RE, Sanderson MW. 2004. Revision of the
Phyllophaga of Hispaniola (Scarabaeidae:
Melolonthinae). Insecta Mundi. 18: 1–154.
Wu W, Cottrell CB, Hansson BS, Löfstedt C. 1999.
Comparative study of pheromone production and
response in Swedish and Zimbabwean populations of
turnip moth, Agrotis segetum.Journal of Chemical
Ecology 25: 177-196.
Zhang A, Robbins PS, Leal WS, Linn CE, Villani MG, Roelofs
WL. 1997. Essential amino acid methyl esters: major sex
pheromone components of the cranberry white grub,
Phyllophaga anxia (Coleoptera: Scarabaeidae). Journal
of Chemical Ecology 23: 231-245.
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Tables
Table 1a.
State or Province City Years Habitat/Trap Placement
Alabama Auburn 97,98,99 sparse pecan grove
Alabama Marion Junction (research farm) 99,00 research farm, edge of woods
British Columbia Aggasiz 99 cranberry bog
Connecticut Vernon 99, 00 suburban
Florida Holiday 98 edge of wooded suburban area
Florida Homestead (research farm) 00 suburban
Georgia Athens 99,00 Athens Botanical Gardens
Georgia Griffin 99 rural farmland/pasture
Georgia Tifton 99,00 rural farmland/pasture
Indiana West Lafayette 2000 suburban - mixed trees and non-maintained turf
Iowa Ames 99,00 grassy area at edge of woods
Kansas Greensburg #1 00 mixed grass range land
Kansas Greensburg #2 01 mixed grass range land
Kansas Manhattan #1 00 golf course
Kansas Manhattan #2 01 golf course
Kentucky Lexington (research farm) 99,00 tall fescue pasture next to tree nursery plantings
Louisiana Chase (research farm) 97, 98 near tree line at the edge of a sweet potato field
Maine Lincolnville Center 99,00 small cranberry bog surrounded by woods
Massachusetts South Amherst 99,00 suburban, edge of woods near horse pasture
Massachusetts Carver 96,98,99,00 large cranberry bog
Massachusetts Lakeville 99,00 cranberry bog near woods
Massachusetts Plympton #1 96 small cranberry bog surrounded by woods
Massachusetts Plympton #2 96 small cranberry bog surrounded by woods
Minnesota St. Paul #1 99 golf course
Minnesota St. Paul #2 00 golf course
Minnesota St. Paul #3 00 golf course
Mississippi Leroy Percy State Park 00 treeline near grass/weed field
Mississippi Sharkey Co., 7 miles SE of Anguilla 00 treeline near grass/weed field
Mississippi Stoneville 00 treeline near grass/weed field
Nebraska Lincoln 98,99,00 suburban
Table 1b.
State or Province City Years Habitat/Trap Placement
New Hampshire Madbury (research farm) 99, 00 pasture grasses surrounded by woods
New Jersey Chatsworth #1 96,98,99 cranberry bog
New Jersey Chatsworth #2 98,99,00 cranberry bog
New Jersey Hammonton 96 commercial blueberry planting
New Jersey New Brunswick 99 edge of woods near horse pasture and vegetables
New York Bellona 98,99,00 edge of woods adjacent to farm land
New York Franklinville 99 edge of woods adjacent to grassy area
New York Riverhead 99 suburban
New York Saratoga Springs 99,00 New York State Tree Nursery
New York Warwick 99,00 golf course
New York Waterloo #1 98,99,00 edge of woods adjacent to grassy area
New York Waterloo #2 98 edge of woods adjacent to grassy area
North Carolina Raleigh #1 (turfgrass research farm) 99 near tall fescue planting
North Carolina Raleigh #2 0 rural subdivision– 50/50 mix of trees and turf
Nova Scotia Kentville Center (research farm) 99,00 edge of woods near pasture, apple orchards
Ohio Columbus (research farm) 0 turfgrass research farm at edge of woods
Ohio Wooster (research farm) 0 edge of woods adjacent to grassy area
Ontario Woodlawn 99,00 edge of cedar/balsam forest near marshy field
Oregon Bandon 99 cranberry bog near woods
Pennsylvania State College (research farm) 99,00 edge of woods adjacent to grassy area
Rhode Island Kingston 99,00 commercial azalea nursery
South Carolina Anderson Co. (research farm). 99 fence line between orchard and pasture near woods
Texas Dallas #1 (research farm) 98,99,00 fence line dividing forage pasture/turf plots
Texas Dallas #2 (research farm) 98 edge of oak tree nursery with weedy undergrowth
Texas Dallas #3 (research farm) 98 edge of mixed wild grasses
Texas Dallas #4 (research farm) 98 edge of mixed wild grasses
Utah Salt Lake City (Red Butte Canyon) 99 grassland/forb community
Vermont Burlington 99,00 grass area near woods
Wisconsin Babcock 99,00 large cranberry bog
Wisconsin Verona (turfgrass research farm) 99,00 wooded fencerow near non-maintained grassy area
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Table 2a. Male catches, sites, and sites-years sorted by species.
Phyllophaga species n sites site-years
aemula 231 1 2
affabilis 77 1 2
anxia TOTAL 20480 33 54
anxia NORTHERN 20460 25 45
anxia SOUTHERN 20 8 9
balia 5 3 4
bipartita 38 2 2
clypeata 2 1 1
congrua 6916 8 10
corrosa 59 2 2
crassissima 5710 11 15
crenulata 15 5 7
crinita 24 7 9
curialis 1 1 1
davisi 6 1 1
diffinis 2 1 1
drakei 91 11 18
ephilida 1726 12 15
fervida 3 1 1
forbesi 141 2 3
forsteri 141 16 26
foxii 1 1 1
fraterna 111 8 14
fraterna-like 37 1 2
fusca 1040 21 37
futilis 7575 20 32
georgiana 2 1 1
glaberrima 70 2 3
glabricula 274 1 2
gracilis 1930 4 7
gracilis var. angulata 6 2 2
hirsuta 2 2 2
hirticula 191 5 7
hirtiventris 1820 7 8
ilicis 1 1 1
implicita 3 2 2
inversa 452 5 8
Table 2b. Male catches, sites, and sites-years sorted by species.
Phyllophaga species n sites site-years
kentuckiana 1 1 1
lanceolata 1 1 1
latifrons 13 1 1
longispina 106 3 4
lota 18 2 2
luctuosa 1 1 1
marginalis 15 5 7
mariana 4 1 2
micans 172 2 2
mississippiensis 1 1 1
nitida 2 2 2
obsoleta 22 1 3
perlonga 2 2 2
postrema 632 4 8
praetermissa 2381 2 2
profunda 1 1 1
quercus 149 7 10
rubiginosa 1594 7 7
rugosa 1393 11 14
soror 11 2 2
submucida 5 1 1
sylvatica 104 2 2
taxodii 2 1 1
torta 4 3 3
ulkei 2 2 2
uniformis 872 1 2
vehemens 79 2 4
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Table 3a. Male catches, sites, and sites-years sorted by n.
Phyllophaga species n sites site-years
anxia TOTAL 20480 33 54
anxia NORTHERN 20460 25 45
futilis 7575 20 32
congrua 6916 8 10
crassissima 5710 11 15
praetermissa 2381 2 2
gracilis 1930 4 7
hirtiventris 1820 7 8
ephilida 1726 12 15
rubiginosa 1594 7 7
rugosa 1393 11 14
fusca 1040 21 37
uniformis 872 1 2
postrema 632 4 8
inversa 452 5 8
glabricula 274 1 2
aemula 231 1 2
hirticula 191 5 7
micans 172 2 2
quercus 149 7 10
forbesi 141 2 3
forsteri 141 16 26
fraterna 111 8 14
longispina 106 3 4
sylvatica 104 2 2
drakei 91 11 18
vehemens 79 2 4
affabilis 77 1 2
glaberrima 70 2 3
corrosa 59 2 2
bipartita 38 2 2
fraterna-like 37 1 2
crinita 24 7 9
obsoleta 22 1 3
anxia SOUTHERN 20 8 9
lota 18 2 2
crenulata 15 5 7
Table 3b. Male catches, sites, and sites-years sorted by n.
Phyllophaga species n sites site-years
marginalis 15 5 7
latifrons 13 1 1
soror 11 2 2
davisi 6 1 1
gracilis var. angulata 6 2 2
balia 5 3 4
submucida 5 1 1
mariana 4 1 2
torta 4 3 3
fervida 3 1 1
implicita 3 2 2
clypeata 2 1 1
diffinis 2 1 1
georgiana 2 1 1
hirsuta 2 2 2
nitida 2 2 2
perlonga 2 2 2
taxodii 2 1 1
ulkei 2 2 2
curialis 1 1 1
foxii 1 1 1
ilicis 1 1 1
kentuckiana 1 1 1
lanceolata 1 1 1
luctuosa 1 1 1
mississippiensis 1 1 1
profunda 1 1 1
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Table 4a. Male catches, sites, and sites-years sorted by sites.
Phyllophaga species n sites site-years
anxia TOTAL 20480 33 54
anxia NORTHERN 20460 25 45
fusca 1040 21 37
futilis 7575 20 32
forsteri 141 16 26
ephilida 1726 12 15
crassissima 5710 11 15
rugosa 1393 11 14
drakei 91 11 18
congrua 6916 8 10
fraterna 111 8 14
anxia SOUTHERN 20 8 9
hirtiventris 1820 7 8
rubiginosa 1594 7 7
quercus 149 7 10
crinita 24 7 9
inversa 452 5 8
hirticula 191 5 7
crenulata 15 5 7
marginalis 15 5 7
gracilis 1930 4 7
postrema 632 4 8
longispina 106 3 4
balia 5 3 4
torta 4 3 3
praetermissa 2381 2 2
micans 172 2 2
forbesi 141 2 3
sylvatica 104 2 2
vehemens 79 2 4
glaberrima 70 2 3
corrosa 59 2 2
bipartita 38 2 2
lota 18 2 2
soror 11 2 2
gracilis var. angulata 6 2 2
implicita 3 2 2
Table 4b. Male catches, sites, and sites-years sorted by sites.
Phyllophaga species n sites site-years
hirsuta 2 2 2
nitida 2 2 2
perlonga 2 2 2
ulkei 2 2 2
uniformis 872 1 2
glabricula 274 1 2
aemula 231 1 2
affabilis 77 1 2
fraterna-like 37 1 2
obsoleta 22 1 3
latifrons 13 1 1
davisi 6 1 1
submucida 5 1 1
mariana 4 1 2
fervida 3 1 1
clypeata 2 1 1
diffinis 2 1 1
georgiana 2 1 1
taxodii 2 1 1
curialis 1 1 1
foxii 1 1 1
ilicis 1 1 1
kentuckiana 1 1 1
lanceolata 1 1 1
luctuosa 1 1 1
mississippiensis 1 1 1
profunda 1 1 1
Journal of Insect Science | www.insectscience.org ISSN: 1536-2442
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Table 5. Alabama, Auburn 1997
Blends indicate the ratio of the methyl esters of L-valine/L-isoleucine
100/0 65/35 50/50 35/65 0/100 BLANK
P. forsteri ♂1 1
P. gracilis ♂14 8 10 13 1068 10
P. hirticula ♂1
P. obsoleta ♂1 2 1 7
P. quercus ♂1 2 4
P.near fraterna ♂1
P. gracilis ♀1 3 2 2 3
P. obsoleta ♀1 1 1 1
P. quercus ♀1 1
A total of 1163 Phyllophaga were taken between 6/9/97 and 8/22/97.
Table 6. Alabama, Auburn 1998
Blends indicate the ratio of the methyl esters of L-valine/L-isoleucine
100/0 95/5 65/35 50/50 35/65 5/95 0/100 BLANK
P. anxia ♂1
P. crinita ♂1
P. diffinis ♂2
P. forsteri ♂2 8
P. gracilis ♂3 9 1 1 14 214 464 4
P. hirticula ♂2 2 26 1
P. obsoleta ♂1 4 2 8 13
P. postrema ♂1
P. praetermissa ♂1 1
P. quercus ♂2 1
P. ulkei ♂1
P. gracilis ♀1 1 1 1 1
P. hirticula ♀1 1
P. obsoleta ♀1 1 1 4 2 1
P. ulkei ♀1
A total of 808 Phyllophaga were taken between 3/30/98 and 8/11/98.
Table 7. Alabama, Auburn 1999
Blends indicate the ratio of the methyl esters of L-valine/L-isoleucine
100/0 90/10 80/20 60/40 40/60 20/80 10/90 0/100 BLANK
P. forsteri ♂4 6 2
P. gracilis ♂2 1 5 2 2 6 19 57 4
P. hirticula ♂1
P. obsoleta ♂1 2 1
P. quercus ♂1 2 4 5
P. gracilis ♀3 1 2
P. obsoleta ♀3 1 1
P. quercus ♀
A total of 140 Phyllophaga were taken between 4/12/99 and 8/17/99.
Table 8. Alabama, Marion Junction 1999
Blends indicate the ratio of the methyl esters of L-valine/L-isoleucine
100/0 90/10 80/20 60/40 40/60 20/80 10/90 0/100 BLANK
P. crenulata ♂3
P. davisi ♂6
P. forbesi ♂121 1
P. futilis ♂1
P. hirtiventris ♂23 2 2
P. micans ♂1
A total of 160 Phyllophaga were taken between 4/5/99 and 7/26/99.
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Table 9. Alabama, Marion Junction 2000
Blends indicate the ratio of the methyl esters of L-valine/L-isoleucine
100/0 90/10 80/20 60/40 40/60 20/80 10/90 0/100 BLANK
P. crinita ♂1 4
P. forbesi ♂10
P. gracilis angulata ♂1
P. hirtiventris ♂19 14 3
P. mississippiensis ♂1
P. crinita ♀1
P. forbesi ♀1 2
P. mississippiensis ♀1
A total of 56 Phyllophaga were taken between 5/19/00 and 8/16/00.
Table 10. Canada, Nova Scotia, Kentville Center 1999
Blends indicate the ratio of the methyl esters of L-valine/L-isoleucine
100/0 90/10 80/20 60/40 40/60 20/80 10/90 0/100 BLANK
P. anxia ♂465 407 385 32 1 3
P. drakei ♂10
P. futilis ♂1
A total of 1304 Phyllophaga were taken between 5/10/99 and 7/2/99.
Table 11. Canada, Nova Scotia, Kentville Center 2000
Blends indicate the ratio of the methyl esters of L-valine/L-isoleucine
100/0 90/10 80/20 60/40 40/60 20/80 10/90 0/100 BLANK
P. anxia ♂668 388 256 25 1
P. drakei ♂32
P. anxia ♀1
A total of 1371 Phyllophaga were taken between 5/22/00 and 7/24/00.
Table 12. Canada, Ontario, Woodlawn 1999
Blends indicate the ratio of the methyl esters of L-valine/L-isoleucine
100/0 90/10 80/20 60/40 40/60 20/80 10/90 0/100 BLANK
P. anxia ♂240 93 35 11 1 1
P. drakei ♂1
P. fusca ♂1 1
P. futilis ♂2 1 1 2 4 6 14 71 1
P. rugosa ♂1 2
A total of 489 Phyllophaga were taken between 5/15/99 and 6/18/99.
Table 13. Canada, Ontario, Woodlawn 2000
Blends indicate the ratio of the methyl esters of L-valine/L-isoleucine
100/0 90/10 80/20 60/40 40/60 20/80 10/90 0/100 BLANK
P. anxia ♂123 80 35 7 3
P. drakei ♂1
P. fusca ♂1 1
P. futilis ♂1 1 1 11 38
A total of 303 Phyllophaga were taken between 5/4/00 and 7/7/00.
Table 14. Connecticut, Vernon 1999
Blends indicate the ratio of the methyl esters of L-valine/L-isoleucine
100/0 90/10 80/20 60/40 40/60 20/80 10/90 0/100 BLANK
P. anxia ♂2 1
P. forsteri ♂2 2 1
P. fraterna ♂2 6 1
P. fusca ♂2 7 6 1
P. hirticula ♂3
A total of 36 Phyllophaga were taken between 5/22/99 and 7/3/99.
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Table 15. Connecticut, Vernon 2000
Blends indicate the ratio of the methyl esters of L-valine/L-isoleucine
100/0 90/10 80/20 60/40 40/60 20/80 10/90 0/100 BLANK
P. forsteri ♂1 1
P. fraterna ♂1
P. fusca ♂1
A total of 4 Phyllophaga were taken on 6/20/00.
Table 16. Florida, Holiday 1998
Blends indicate the ratio of the methyl esters of L-valine/L-isoleucine
100/0 95/5 65/35 50/50 35/65 5/95 0/100 BLANK
P. glaberrima ♂6 14 4 12 3
P. latifrons ♂11 2
A total of 52 Phyllophaga were taken between 4/22/98 and 7/16/98.
Table 17. Georgia, Athens 1999
Blends indicate the ratio of the methyl esters of L-valine/L-isoleucine
100/0 90/10 80/20 60/40 40/60 20/80 10/90 0/100 BLANK
P. aemula ♂16 48 18 1 1 1
P. ephilida ♂6
P. forsteri ♂1
A total of 93 Phyllophaga were taken between 5/28/99 and 9/20/99.
Table 18. Georgia, Athens 2000
Blends indicate the ratio of the methyl esters of L-valine/L-isoleucine
100/0 90/10 80/20 60/40 40/60 20/80 10/90 0/100 BLANK
P. aemula ♂21 96 23 4 1
P. anxia ♂1
P. ephilida ♂20 1
P. fusca ♂2
P. curialis ♂1
P. quercus ♂1 9 2 17 2 1
A total of 202 Phyllophaga were taken between 5/11/00 and 9/14/00.
Table 19. Georgia, Griffin 1999
Blends indicate the ratio of the methyl esters of L-valine/L-isoleucine
100/0 90/10 80/20 60/40 40/60 20/80 10/90 0/100 BLANK
P. anxia ♂1
P. ephilida ♂1
P. forsteri ♂5 3
P. ilicis ♂1
P. quercus ♂1
A total of 12 Phyllophaga were taken between 4/22/99 and 7/18/99.
Table 20. Georgia, Tifton 1999
Blends indicate the ratio of the methyl esters of L-valine/L-isoleucine
100/0 90/10 80/20 60/40 40/60 20/80 10/90 0/100 BLANK
P. glaberrima ♂4 3
P. luctuosa ♂1
P. mariana ♂1 2
P. postrema ♂3
P. quercus ♂1 2
P. ulkei ♂1
P. uniformis ♂179 135 108 18
A total of 458 Phyllophaga were taken between 4/16/99 and 7/30/99.
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Table 21. Georgia, Tifton 2000
Blends indicate the ratio of the methyl esters of L-valine/L-isoleucine
100/0 90/10 80/20 60/40 40/60 20/80 10/90 0/100 BLANK
P. clypeata ♂1 1
P. georgiana ♂2
P. glaberrima ♂15 7 2
P. lota ♂1 3 1
P. mariana ♂1
P. quercus ♂2
P. uniformis ♂2 100 205 118 6 1 1
P. uniformis ♀1 1 1 1
A total of 473 Phyllophaga were taken between 5/22/00 and 9/1/00.
Table 22. Iowa, Ames, East Reactor Woods 1999
Blends indicate the ratio of the methyl esters of L-valine/L-isoleucine
100/0 90/10 80/20 60/40 40/60 20/80 10/90 0/100 BLANK
P. anxia ♂1
P. futilis ♂1 1 1 1 60
A total of 65 Phyllophaga were taken between 5/19/99 and 6/28/99.
Table 23. Iowa, Ames, East Reactor Woods 2000
Blends indicate the ratio of the methyl esters of L-valine/L-isoleucine
100/0 90/10 80/20 60/40 40/60 20/80 10/90 0/100 BLANK
P. anxia ♂1
P. futilis ♂12
P. rugosa ♂1
P. futilis ♀1
A total of 15 Phyllophaga were taken between 6/5/00 and 7/10/00.
Table 24. Indiana, West Lafayette 2000
Blends indicate the ratio of the methyl esters of L-valine/L-isoleucine
100/0 90/10 80/20 60/40 40/60 20/80 10/90 0/100 BLANK
P. balia ♂2
P. ephilida ♂5
P. fraterna ♂1
P. futilis ♂1 4 55
A total of 68 Phyllophaga were taken between 5/30/00 and 8/17/00.
Table 25. Kansas, Greensburg #1 2000
Blends indicate the ratio of the methyl esters of L-valine/L-isoleucine
100/0 90/10 80/20 60/40 40/60 20/80 10/90 0/100 BLANK
P. corrosa ♂13 24 17 4
P. crassissima ♂5 7 2 14 2
P. gracilis angulata ♂5
P. lanceolata ♂1
P. rubiginosa ♂1 6 5 1
P. submucida ♂3 1 1
A total of 112 Phyllophaga were taken between 4/29/00 and 7/30/00.
Table 26. Kansas, Greensburg #2 2001
Blends indicate the ratio of the methyl esters of L-valine/L-isoleucine
100/0 90/10 80/20 60/40 40/60 20/80 10/90 0/100 BLANK
P. congrua ♂211 33
P. corrosa ♂1
P. crassissima ♂1 6 7 23 189 640 1593 277 7
P. praetermissa ♂35 583 945 693 100 7 9 7 11
P. rubiginosa ♂1 18 5 11 2 11
P. crassissima ♀1 4
P. rubiginosa ♀2 1
A total of 5434 Phyllophaga were taken between 4/29/01 and 6/20/01.
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Table 27. Kansas, Manhattan #1 2000
Blends indicate the ratio of the methyl esters of L-valine/L-isoleucine
100/0 90/10 80/20 60/40 40/60 20/80 10/90 0/100 BLANK
P. affabilis ♂7 29 22
P. bipartita ♂1 1 13
P. crassissima ♂1 4 15 56 96 90
P. ephilida ♂28
P. fusca ♂2 5
P. futilis ♂3 1 50
P. glabricula ♂1 41 112 34 10
P. implicita ♂1
P. inversa ♂155 13 2
P. rubiginosa ♂1 30 43 66 18 4
P. sylvatica ♂1 1 4 1
A total of 962 Phyllophaga were taken between 5/4/00 and 8/22/00.
Table 28. Kansas, Manhattan #2 2001
Blends indicate the ratio of the methyl esters of L-valine/L-isoleucine
100/0 90/10 80/20 60/40 40/60 20/80 10/90 0/100 BLANK
P. affabilis ♂7 10 2
P. bipartita ♂23
P. crassissima ♂1 15 71 65
P. ephilida ♂28
P. fusca ♂1 4 1 11 23 63 101 11
P. futilis ♂1 3 47
P. glabricula ♂8 34 31 2 1
P. inversa ♂153 21
P. rubiginosa ♂5 143 389 295 405 107 21 1
P. rugosa ♂1
P. sylvatica ♂18 25 39 13 2
P. vehemens ♂29 7 2 1 3
A total of 2245 Phyllophaga were taken between 4/30/01 and 8/16/01.
Table 29. Kentucky, Lexington 1999
Blends indicate the ratio of the methyl esters of L-valine/L-isoleucine
100/0 90/10 80/20 60/40 40/60 20/80 10/90 0/100 BLANK
P. ephilida ♂4 2 1 3 1 17 3 455 5
P. fusca ♂3 5 10 14 8 5
P. futilis ♂2 1 3 3 15 72 445 1
P. hirticula ♂1 3
P. inversa ♂15 4 1
P. rugosa ♂1 32 64 17 2 2 3
P. ephilida ♀2 1 1
P. futilis ♀1 1
P. rugosa ♀2
A total of 1231 Phyllophaga were taken between 4/22/99 and 8/20/99.
Table 30. Kentucky, Lexington 2000
Blends indicate the ratio of the methyl esters of L-valine/L-isoleucine
100/0 90/10 80/20 60/40 40/60 20/80 10/90 0/100 BLANK
P. ephilida ♂11 5 5 1 2 3 9 994 2
P. fusca ♂2 5 2 3 11 9
P. futilis ♂1 10 5 7 2 247 209 1359 6
P. hirticula ♂1 2 5 146 4
P. inversa ♂65 2
P. rugosa ♂17 593 504 76 8 3 1 1 2
P. futilis ♀1
P. rugosa ♀3 1
A total of 4347 Phyllophaga were taken between 5/2/00 and 8/11/00.
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Table 31. Louisiana, Chase 1997
Blends indicate the ratio of the methyl esters of L-valine/L-isoleucine
100/0 65/35 50/50 35/65 0/100 BLANK
P. ephilida ♂1 1
P. forbesi ♂10
P. quercus ♂15 15 9
P. taxodii ♂2
A total of 53 Phyllophaga were taken between 6/17/97 and 8/15/97.
Table 32. Massachusetts, South Amherst 1999
Blends indicate the ratio of the methyl esters of L-valine/L-isoleucine
100/0 90/10 80/20 60/40 40/60 20/80 10/90 0/100 BLANK
P. anxia ♂4 6 2 2 3 291
P. drakei ♂1
P. forsteri ♂7 12 4 1
P. fraterna ♂12 16 5 1
P. fusca ♂2 14 68 62 59 7 1
P. gracilis ♂1
P. longispina ♂57
P. anxia ♀1 1
A total of 642 Phyllophaga were taken between 5/7/99 and 7/20/99.
Table 33. Massachusetts, South Amherst 2000
Blends indicate the ratio of the methyl esters of L-valine/L-isoleucine
100/0 90/10 80/20 60/40 40/60 20/80 10/90 0/100 BLANK
P. anxia ♂4 1 1 76
P. forsteri ♂1 3 1
P. fraterna ♂2 6
P. fusca ♂1 1 2 5 5 1
P. gracilis ♂1
P. longispina ♂1 45
A total of 157 Phyllophaga were taken between 5/8/00 and 7/28/00.
Table 34. Massachusetts, Carver 1996
Blends indicate the ratio of the methyl esters of L-valine/L-isoleucine
100/0 65/35 50/50 35/65 0/100 BLANK
P. anxia ♂18 700 591 515 7 3
P. drakei ♂1
P. anxia ♀1 3
A total of 1839 Phyllophaga were taken between 6/4/96 and 6/10/96.
Table 35. Massachusetts, Carver 1998
Blends indicate the ratio of the methyl esters of L-valine/L-isoleucine
100/0 95/5 65/35 50/50 35/65 5/95 0/100 BLANK
P. anxia ♂13 67 627 682 608 54 11 10
P. drakei ♂1
P. forsteri ♂4
P. anxia ♀1 1 2 1
A total of 2082 Phyllophaga were taken between 5/22/98 and 7/7/98.
Table 36. Massachusetts, Carver 1999
Blends indicate the ratio of the methyl esters of L-valine/L-isoleucine
100/0 90/10 80/20 60/40 40/60 20/80 10/90 0/100 BLANK
P. anxia ♂1 115 302 440 514 232 88 2 2
P. crenulata ♂1
P. drakei ♂1
P. anxia ♀1 1 1 2
A total of 1703 Phyllophaga were taken between 5/20/99 and 7/12/99.
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Table 37. Massachusetts, Carver 2000
Blends indicate the ratio of the methyl esters of L-valine/L-isoleucine
100/0 90/10 80/20 60/40 40/60 20/80 10/90 0/100 BLANK
P. anxia ♂1 127 318 365 356 169 79 2 2
P. drakei ♂2
P. forsteri ♂1
P. hirsuta ♂1
P. anxia ♀1 1 3
A total of 1428 Phyllophaga were taken between 5/15/00 and 7/10/00.
Table 38. Massachusetts, Lakeville 1999
Blends indicate the ratio of the methyl esters of L-valine/L-isoleucine
100/0 90/10 80/20 60/40 40/60 20/80 10/90 0/100 BLANK
P. anxia ♂77 114 97 38 26 9 2
P. drakei ♂1
P. forsteri ♂8 4 2
P. fusca ♂2 3
P. gracilis ♂1
P. hirsuta ♂1
P. marginalis ♂1
P. crenulata ♀1
A total of 390 Phyllophaga were taken between 5/10/99 and 7/12/99.
Table 39. Massachusetts, Lakeville 2000
Blends indicate the ratio of the methyl esters of L-valine/L-isoleucine
100/0 90/10 80/20 60/40 40/60 20/80 10/90 0/100 BLANK
P. anxia ♂23 22 27 12 2 1
P. drakei ♂2
P. forsteri ♂15 4 1
P. fusca ♂1
P. marginalis ♂2
A total of 112 Phyllophaga were taken between 5/2/00 and 7/12/00.
Table 40. Massachusetts, Plympton #1 1996
Blends indicate the ratio of the methyl esters of L-valine/L-isoleucine
100/0 65/35 50/50 35/65 0/100 BLANK
P. anxia ♂8 151 128 105
P. drakei ♂1
P. forsteri ♂3
P. fraterna ♂1
P. anxia ♀1
A total of 392 Phyllophaga were taken between 6/4/96 and 6/10/96.
Table 41. Massachusetts, Plympton #2 1996
Blends indicate the ratio of the methyl esters of L-valine/L-isoleucine
100/0 65/35 50/50 35/65 0/100 BLANK
P. anxia ♂12 146 128 110 2 3
P. drakei ♂2
P. forsteri ♂2
P. fusca ♂1
A total of 406 Phyllophaga were taken between 6/4/96 and 6/10/96.
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Journal of Insect Science: Vol. 6 | Article 39 113

Table 42. Maine, Lincolnville Center 1999
Blends indicate the ratio of the methyl esters of L-valine/L-isoleucine
100/0 90/10 80/20 60/40 40/60 20/80 10/90 0/100 BLANK
P. anxia ♂327 169 92 16 5 2 4 119
P. drakei ♂2
P. fusca ♂2 2 9 7 2
P. fraterna ♂1 8 1 4 1
P. anxia ♀1
A total of 774 Phyllophaga were taken between 5/18/99 and 7/2/99.
Table 43. Maine, Lincolnville Center 2000
Blends indicate the ratio of the methyl esters of L-valine/L-isoleucine
100/0 90/10 80/20 60/40 40/60 20/80 10/90 0/100 BLANK
P. anxia ♂665 272 152 52 6 2 8 128
P. drakei ♂1
P. fusca ♂1 2 3 4 5
P. fraterna ♂1 4 1
P. marginalis ♂1
P. anxia ♀1
A total of 1310 Phyllophaga were taken between 5/22/00 and 8/21/00.
Table 44. Minnesota, St. Paul #1 1999
Blends indicate the ratio of the methyl esters of L-valine/L-isoleucine
100/0 90/10 80/20 60/40 40/60 20/80 10/90 0/100 BLANK
P. anxia ♂2
P. futilis ♂1 1 31
P. rugosa ♂1
A total of 36 Phyllophaga were taken between 6/7/99 and 6/13/99.
Table 45. Minnesota, St. Paul #1 2000
Blends indicate the ratio of the methyl esters of L-valine/L-isoleucine
100/0 90/10 80/20 60/40 40/60 20/80 10/90 0/100 BLANK
P. anxia ♂2 1
P. futilis ♂1 4 2 103
P. rugosa ♂2 1 1
P. futilis ♀1 1 1
P. rugosa ♀2 2 1
A total of 125 Phyllophaga were taken between 6/9/00 and 6/30/00.
Table 46. Minnesota, St. Paul #2 2000
Blends indicate the ratio of the methyl esters of L-valine/L-isoleucine
100/0 90/10 80/20 60/40 40/60 20/80 10/90 0/100 BLANK
P. anxia ♂5
P. futilis ♂1 6
P. nitida ♂1
P. rugosa ♂2
A total of 15 Phyllophaga were taken between 6/9/00 and 6/30/00.
Table 47. Mississippi, Leroy Percy State Park 2000
Blends indicate the ratio of the methyl esters of L-valine/L-isoleucine
100/0 90/10 80/20 60/40 40/60 20/80 10/90 0/100 BLANK
P. congrua ♂24 15 7 4 1 1
P. crassissima ♂1 2
P. hirtiventris ♂441 206 69 7 4 1 1 2 2
P. congrua ♀2 2 1
P. hirtiventris ♀1
A total of 794 Phyllophaga were taken between 5/5/00 and 6/26/00.
Journal of Insect Science | www.insectscience.org ISSN: 1536-2442
Journal of Insect Science: Vol. 6 | Article 39 114

Table 48. Mississippi, Sharkey County, 5 miles SE of Anguilla 2000
Blends indicate the ratio of the methyl esters of L-valine/L-isoleucine
100/0 90/10 80/20 60/40 40/60 20/80 10/90 0/100 BLANK
P. congrua ♂213 158 146 48 21 2 1 1
P. hirtiventris ♂24 10 5 1
P. perlonga ♂1
P. profunda ♂1
P. congrua ♀1 1
A total of 667 Phyllophaga were taken between 5/12/00 and 6/22/00.
Table 49. Mississippi, Stoneville 2000
Blends indicate the ratio of the methyl esters of L-valine/L-isoleucine
100/0 90/10 80/20 60/40 40/60 20/80 10/90 0/100 BLANK
P. anxia ♂1
P. congrua ♂114 116 69 30 4 8
P. crassissima ♂2 21 26 2
P. ephilida ♂1 1
P. hirtiventris ♂442 324 183 19 10 1 1
P. perlonga ♂1
P. congrua ♀1 1 3 6 1 2 1
P.hirtiventris ♀1 1 1 1
A total of 1395 Phyllophaga were taken between 5/3/00 and 7/10/00.
Table 50. Nebraska, Lincoln 1998
Blends indicate the ratio of the methyl esters of L-valine/L-isoleucine
100/0 95/5 65/35 50/50 35/65 5/95 0/100 BLANK
P. crassissima ♂25 19 44 24 1
P. crenulata ♂1
P. futilis ♂1 1 1 31 355
P. inversa ♂1
P. vehemens ♂1
A total of 569 Phyllophaga were taken between 5/15/98 and 6/29/98.
Table 51. Nebraska, Lincoln 1999
Blends indicate the ratio of the methyl esters of L-valine/L-isoleucine
100/0 90/10 80/20 60/40 40/60 20/80 10/90 0/100 BLANK
P. crassissima ♂5 23 82 113 87
P. crenulata ♂5
P. fusca ♂2 3
P. futilis ♂1 4 9 11 11 20 924
P. implicita ♂2
P. vehemens ♂31 1
P. futilis ♀1
P. implicita ♀1 1
A total of 1337 Phyllophaga were taken between 5/16/99 and 7/10/99.
Table 52. Nebraska, Lincoln 2000
Blends indicate the ratio of the methyl esters of L-valine/L-isoleucine
100/0 90/10 80/20 60/40 40/60 20/80 10/90 0/100 BLANK
P. crassissima ♂1 16 76 41 38
P. crenulata ♂1
P. fusca ♂1 2
P. futilis ♂2 2 3 6 18 489
P. rugosa ♂1
P. vehemens ♂3 1
P. futilis ♀1
P. implicita ♀1
A total of 703 Phyllophaga were taken between 5/10/00 and 7/3/00.
Journal of Insect Science | www.insectscience.org ISSN: 1536-2442
Journal of Insect Science: Vol. 6 | Article 39 115

Table 53. New Hampshire, Madbury 1999
Blends indicate the ratio of the methyl esters of L-valine/L-isoleucine
100/0 90/10 80/20 60/40 40/60 20/80 10/90 0/100 BLANK
P. anxia♂6 2 2 139
P. forsteri ♂1
P. fraterna ♂5 2
P. fusca♂1 1 3 2 2 1
P. gracilis ♂2 12
P. anxia ♀1
P. fraterna ♀1 1 1
P. hirticula ♀1
A total of 186 Phyllophaga were taken between 5/9/99 and 8/3/99.
Table 54. New Hampshire, Madbury 2000
Blends indicate the ratio of the methyl esters of L-valine/L-isoleucine
100/0 90/10 80/20 60/40 40/60 20/80 10/90 0/100 BLANK
P. anxia♂7 3 30
P. forsteri ♂1
P. fraterna ♂1 2 2
P. fusca♂1 2 1 4
P. lonispina ♂2
P. fusca ♀1
A total of 57 Phyllophaga were taken between 5/8/00 and 7/17/00.
Table 55. New Jersey, Chatsworth #1 1996
Blends indicate the ratio of the methyl esters of L-valine/L-isoleucine
100/0 65/35 50/50 35/65 0/100 BLANK
P. anxia ♂2
P. forsteri ♂1
P. postrema ♂1 61
A total of 65 Phyllophaga were taken between 5/20/96 and 7/1/96.
Table 56. New Jersey, Chatsworth #1 1998
Blends indicate the ratio of the methyl esters of L-valine/L-isoleucine
100/0 95/5 65/35 50/50 35/65 5/95 0/100 BLANK
P. forsteri ♂1
P. postrema ♀1 10 32
A total of 44 Phyllophaga were taken between 5/15/98 and 6/26/98.
Table 57. New Jersey, Chatsworth #1 1999
Blends indicate the ratio of the methyl esters of L-valine/L-isoleucine
100/0 90/10 80/20 60/40 40/60 20/80 10/90 0/100 BLANK
P. forsteri ♂1
P. postrema ♂1 1 1 1 73
A total of 77 Phyllophaga were taken between 5/24/99 and 7/13/99.
Table 58. New Jersey, Chatsworth #2 1998
Blends indicate the ratio of the methyl esters of L-valine/L-isoleucine
100/0 95/5 65/35 50/50 35/65 5/95 0/100 BLANK
P. anxia ♂1
P. postrema ♂1 1 1 1 54 128
P. postrema ♀1
A total of 188 Phyllophaga were taken between 5/15/98 and 6/26/98.
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Table 59. New Jersey, Chatsworth #2 1999
Blends indicate the ratio of the methyl esters of L-valine/L-isoleucine
100/0 90/10 80/20 60/40 40/60 20/80 10/90 0/100 BLANK
P. postrema ♂1 1 2 77
P. postrema ♀1 1
A total of 83 Phyllophaga were taken between 6/11/99 and 7/19/99.
Table 60. New Jersey, Chatsworth #2 2000
Blends indicate the ratio of the methyl esters of L-valine/L-isoleucine
100/0 90/10 80/20 60/40 40/60 20/80 10/90 0/100 BLANK
P. anxia ♂2
P. forsteri ♂1 3 1
P. postrema ♂1 6 3 169
P. postrema ♀1
A total of 188 Phyllophaga were taken between 5/18/00 and 7/4/00.
Table 61. New Jersey, Hammonton 1996
Blends indicate the ratio of the methyl esters of L-valine/L-isoleucine
100/0 65/35 50/50 35/65 0/100 BLANK
P. anxia ♂11
P. forsteri ♂3
A total of 14 Phyllophaga were taken between 5/11/96 and 7/1/96.
Table 62. New Jersey, New Brunswick 1999
Blends indicate the ratio of the methyl esters of L-valine/L-isoleucine
100/0 90/10 80/20 60/40 40/60 20/80 10/90 0/100 BLANK
P. crenulata ♂1
P. ephilida ♂2
P. futilis ♂2 2 1
A total of 8 Phyllophaga were taken between 6/1/99 and 8/10/99.
Table 63. New York, Bellona 1998
Blends indicate the ratio of the methyl esters of L-valine/L-isoleucine
100/0 95/5 65/35 50/50 35/65 5/95 0/100 BLANK
P. anxia ♂171 76 10 6 2 4 1
P. fusca ♂2 5 7
P. futilis ♂1
P. hirticula ♂1
A total of 286 Phyllophaga were taken between 5/13/98 and 6/26/98.
Table 64. New York, Bellona 1999
Blends indicate the ratio of the methyl esters of L-valine/L-isoleucine
100/0 90/10 80/20 60/40 40/60 20/80 10/90 0/100 BLANK
P. anxia ♂128 28 4 3
P. fusca ♂1 3 4
P. futilis ♂3
A total of 181 Phyllophaga were taken between 5/5/99 and 6/18/99.
Table 65. New York, Bellona 2000
Blends indicate the ratio of the methyl esters of L-valine/L-isoleucine
100/0 90/10 80/20 60/40 40/60 20/80 10/90 0/100 BLANK
P. anxia ♂61 19 6 1
P. balia ♂1
P. futilis ♂1 2
A total of 91 Phyllophaga were taken between 5/10/00 and 6/12/00.
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Table 66. New York, Franklinville 1999
Blends indicate the ratio of the methyl esters of L-valine/L-isoleucine
100/0 90/10 80/20 60/40 40/60 20/80 10/90 0/100 BLANK
P. anxia ♂282 147 25 2 2 1 3 122
P. fusca ♂1 1
P. futilis ♂1
P. marginalis ♂3
P. fusca ♀1
A total of 591 Phyllophaga were taken between 5/1/99 and 6/13/99.
Table 67. New York, Riverhead 1999
Blends indicate the ratio of the methyl esters of L-valine/L-isoleucine
100/0 90/10 80/20 60/40 40/60 20/80 10/90 0/100 BLANK
P. forsteri ♂1
P. fusca ♂1
A total of 2 Phyllophaga were taken between 4/23/99 and 7/16/99.
Table 68. New York, Saratoga Springs 1999
Blends indicate the ratio of the methyl esters of L-valine/L-isoleucine
100/0 90/10 80/20 60/40 40/60 20/80 10/90 0/100 BLANK
P. anxia ♂5 1 1
P. fusca ♂1
A total of 8 Phyllophaga were taken between 5/10/99 and 6/4/99.
Table 69. New York, Saratoga Springs 2000
Blends indicate the ratio of the methyl esters of L-valine/L-isoleucine
100/0 90/10 80/20 60/40 40/60 20/80 10/90 0/100 BLANK
P. anxia ♂1 1
P. crenulata ♂2 1 2
P. crenulata ♀2
A total of 9 Phyllophaga were taken between 5/19/00 and 6/21/00.
Table 70. New York, Warwick 1999
Blends indicate the ratio of the methyl esters of L-valine/L-isoleucine
100/0 90/10 80/20 60/40 40/60 20/80 10/90 0/100 BLANK
P. anxia ♂48 17 5
P. balia ♂1
P. fusca ♂13 45 14 20
P. marginalis ♂1
A total of 164 Phyllophaga were taken between 5/10/99 and 6/10/99.
Table 71. New York, Warwick 2000
Blends indicate the ratio of the methyl esters of L-valine/L-isoleucine
100/0 90/10 80/20 60/40 40/60 20/80 10/90 0/100 BLANK
P. anxia ♂66 23 7
P. balia ♂1
P. drakei ♂21
P. fusca ♂1 7 11 25 3
P. marginalis ♂2
A total of 167 Phyllophaga were taken between 5/11/00 and 7/20/00.
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Table 72. New York, Waterloo #1 1998
Blends indicate the ratio of the methyl esters of L-valine/L-isoleucine
100/0 95/5 65/35 50/50 35/65 5/95 0/100 BLANK
P. anxia ♂299 140 4 3 1 7 33
P. fusca ♂1
P. futilis ♂1 2 3 2 46 191
P. futilis ♀1
A total of 734 Phyllophaga were taken between 5/15/98 and 7/2/98.
Table 73. New York, Waterloo #1 1999
Blends indicate the ratio of the methyl esters of L-valine/L-isoleucine
100/0 90/10 80/20 60/40 40/60 20/80 10/90 0/100 BLANK
P. anxia ♂695 286 40 5 3 1 16 60
P. fusca ♂1 4 7 4 1
P. futilis ♂1 1 1 2 2 2 26 165
A total of 1323 Phyllophaga were taken between 5/10/99 and 6/18/99.
Table 74. New York, Waterloo #1 2000
Blends indicate the ratio of the methyl esters of L-valine/L-isoleucine
100/0 90/10 80/20 60/40 40/60 20/80 10/90 0/100 BLANK
P. anxia ♂404 112 15 1 2 4 17
P. fusca ♂2
P. futilis ♂1 2 3 9 17 65 309
A total of 963 Phyllophaga were taken between 5/8/00 and 6/19/00.
Table 75. New York, Waterloo #2 1998
Blends indicate the ratio of the methyl esters of L-valine/L-isoleucine
100/0 95/5 65/35 50/50 35/65 5/95 0/100 BLANK
P. anxia ♂84 36 6 1 3 6
P. futilis ♂6 1 15 2 65 78
P. futilis ♀1 1 1
A total of 306 Phyllophaga were taken between 5/15/98 and 6/26/98.
Table 76. North Carolina, Raleigh #1 1999
Blends indicate the ratio of the methyl esters of L-valine/L-isoleucine
100/0 90/10 80/20 60/40 40/60 20/80 10/90 0/100 BLANK
P. ephilida ♂1 61
P. soror ♂1
P. ephilida ♀1
A total of 64 Phyllophaga were taken between 7/6/99 and 7/26/99.
Table 77. North Carolina, Raleigh #2 2000
Blends indicate the ratio of the methyl esters of L-valine/L-isoleucine
100/0 90/10 80/20 60/40 40/60 20/80 10/90 0/100 BLANK
P. anxia ♂2
P. ephilida ♂1 33 11
P. forsteri ♂1 4 2
P. foxii ♂1
P. hirsuta ♂1
P. micans ♂5 1 5 19 29 111 1
P. quercus ♂5 6 32 5
P. soror ♂10
P. ephilida ♀1
P. micans ♀3 2 2
P. soror ♀1
A total of 297 Phyllophaga were taken between 5/3/00 and 9/5/00.
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Table 78. Ohio, Columbus 2000
Blends indicate the ratio of the methyl esters of L-valine/L-isoleucine
100/0 90/10 80/20 60/40 40/60 20/80 10/90 0/100 BLANK
P. anxia ♂1
P. futilis ♂1 2 2 4 34 212
P. inversa ♂7 1 1
P. kentuckiana ♂1
P. futilis ♀1
A total of 267 Phyllophaga were taken between 5/3/00 and 6/28/00.
Table 79. Ohio, Wooster 2000
Blends indicate the ratio of the methyl esters of L-valine/L-isoleucine
100/0 90/10 80/20 60/40 40/60 20/80 10/90 0/100 BLANK
P. futilis ♂1
P. longispina ♂1
P. lota ♂4 3 6
P. rugosa ♂1 2
A total of 18 Phyllophaga were taken between 5/13/00 and 6/20/00.
Table 80. Pennsylvania, State College 1999
Blends indicate the ratio of the methyl esters of L-valine/L-isoleucine
100/0 90/10 80/20 60/40 40/60 20/80 10/90 0/100 BLANK
P. anxia ♂3 1
P. fusca ♂4 4 34 60 71 17
P. futilis ♂1 12 333
P. fraterna ♂1 2 3 3 3 1
P. fraterna-like ♂18
P. inversa ♂9
P. fusca ♀1
P. futilis ♀2 1
P. fraterna ♀1
A total of 585 Phyllophaga were taken between 5/7/99 and 7/2/99.
Table 81. Pennsylvania, State College 2000
Blends indicate the ratio of the methyl esters of L-valine/L-isoleucine
100/0 90/10 80/20 60/40 40/60 20/80 10/90 0/100 BLANK
P. anxia ♂2 1
P. fusca ♂4 4 3 1
P. futilis ♂1 1 3 4 92 445 1
P. fraterna ♂2 3
P. fraterna-like ♂19
P. inversa ♂1
P. futilis ♀1
A total of 588 Phyllophaga were taken between 5/1/00 and 6/28/00.
Table 82. Rhode Island, Kingston 1999
Blends indicate the ratio of the methyl esters of L-valine/L-isoleucine
100/0 90/10 80/20 60/40 40/60 20/80 10/90 0/100 BLANK
P. anxia ♂136 449 512 336 218 106 31 17 14
P. forsteri ♂1 4 1
P. fusca ♂1
P. anxia ♀1 2 2 4 4 2 3 3 3
A total of 1850 Phyllophaga were taken between 5/14/99 and 6/25/99.
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Table 83. Rhode Island, Kingston 2000
Blends indicate the ratio of the methyl esters of L-valine/L-isoleucine
100/0 90/10 80/20 60/40 40/60 20/80 10/90 0/100 BLANK
P. anxia ♂14 58 59 42 16 1 2 20 1
P. forsteri ♂1
P. marginalis ♂5
P. anxia ♀1 3
A total of 224 Phyllophaga were taken between 5/15/00 and 7/5/00.
Table 84. South Carolina, Anderson County 1999
Blends indicate the ratio of the methyl esters of L-valine/L-isoleucine
100/0 90/10 80/20 60/40 40/60 20/80 10/90 0/100 BLANK
P. ephilida ♂2 13
P. fervida ♂3
P. hirticula ♂1 1
P. quercus ♂1
P. fervida ♀1
A total of 22 Phyllophaga were taken between 5/12/99 and 8/20/99.
Table 85. Texas, Dallas #1 1998
Blends indicate the ratio of the methyl esters of L-valine/L-isoleucine
100/0 95/5 65/35 50/50 35/65 5/95 0/100 BLANK
P. congrua ♂531 307 42 16 5 3 3
P. crassissima ♂81 147 426 137 6 1
P. crinita ♂1 1
P. hirtiventris ♂1
P. torta ♂1
P. crassissima ♀2
P. torta ♀1
A total of 1712 Phyllophaga were taken between 4/22/98 and 6/15/98.
Table 86. Texas, Dallas #1 1999
Blends indicate the ratio of the methyl esters of L-valine/L-isoleucine
100/0 90/10 80/20 60/40 40/60 20/80 10/90 0/100 BLANK
P. congrua ♂42 30 58
P. crassissima ♂5 14 10
A total of 159 Phyllophaga were taken between 5/13/99 and 5/19/99.
Table 87. Texas, Dallas #1 2000
Blends indicate the ratio of the methyl esters of L-valine/L-isoleucine
100/0 90/10 80/20 60/40 40/60 20/80 10/90 0/100 BLANK
P. congrua ♂304 168 99 16 5 2 1
P. crassissima ♂2 1 5 61 81 205 76 12 1
P. crinita ♂3 1 1 2 2
P. torta ♂1 1
P. crassissima ♀1 1
A total of 1053 Phyllophaga were taken between 4/28/00 and 6/27/00.
Table 88. Texas, Dallas #2 1998
Blends indicate the ratio of the methyl esters of L-valine/L-isoleucine
100/0 95/5 65/35 50/50 35/65 5/95 0/100 BLANK
P. congrua ♂760 489 45 22 4
P. crassissima ♂10 18 59 22 1
P. crinita ♂1 2 1
P. hirtiventris ♂1 1
P. rubiginosa ♂1 1
P. congrua ♀2 1
P. crinita ♀1 1
A total of 1443 Phyllophaga were taken between 4/22/98 and 6/15/98.
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Table 89. Texas, Dallas #3 1998
Blends indicate the ratio of the methyl esters of L-valine/L-isoleucine
100/0 95/5 65/35 50/50 35/65 5/95 0/100 BLANK
P. congrua ♂654 711 28 17 2 4
P. crassissima ♂6 22 95 28 1
P. crinita ♂1
P. hirtiventris ♂3 1
P. rubiginosa ♂1 1
P. torta ♂1
A total of 1576 Phyllophaga were taken between 4/22/98 and 6/15/98.
Table 90. Texas, Dallas #4 1998
Blends indicate the ratio of the methyl esters of L-valine/L-isoleucine
100/0 95/5 65/35 50/50 35/65 5/95 0/100 BLANK
P. congrua ♂533 694 65 27 10 1 2
P. crassissima ♂30 86 183 44 2
P. crinita ♂1 1
P. rubiginosa ♂1
P. congrua ♀1
A total of 1681 Phyllophaga were taken between 4/22/98 and 6/15/98.
Table 91. Utah, Salt Lake City 1999
Blends indicate the ratio of the methyl esters of L-valine/L-isoleucine
100/0 90/10 80/20 60/40 40/60 20/80 10/90 0/100 BLANK
P. drakei ♂2
A total of 2 Phyllophaga were taken between 6/2/99 and 7/30/99.
Table 92. Vermont, Burlington 1999
Blends indicate the ratio of the methyl esters of L-valine/L-isoleucine
100/0 90/10 80/20 60/40 40/60 20/80 10/90 0/100 BLANK
P. anxia ♂12 7 5 1 1 1 30
P. fraterna ♂2 2
P. fusca ♂2 3 2 7 4 3
A total of 83 Phyllophaga were taken between 5/7/99 and 6/20/99.
Table 93. Vermont, Burlington 2000
Blends indicate the ratio of the methyl esters of L-valine/L-isoleucine
100/0 90/10 80/20 60/40 40/60 20/80 10/90 0/100 BLANK
P. anxia ♂3 9 1
P. fraterna ♂1 1
P. fusca ♂4 2 2
A total of 23 Phyllophaga were taken between 5/21/00 and 7/5/00.
Table 94. Wisconsin, Babcock 1999
Blends indicate the ratio of the methyl esters of L-valine/L-isoleucine
100/0 90/10 80/20 60/40 40/60 20/80 10/90 0/100 BLANK
P. anxia ♂181 30 6 8 7 3 8 12 2
P. rugosa ♂2 1 1
P. anxia ♀1
A total of 262 Phyllophaga were taken between 5/7/99 and 6/29/99.
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Table 95. Wisconsin, Babcock 2000
Blends indicate the ratio of the methyl esters of L-valine/L-isoleucine
100/0 90/10 80/20 60/40 40/60 20/80 10/90 0/100 BLANK
P. anxia ♂259 63 19 2 4 12 25 20 1
P. drakei ♂9 3
P. rugosa ♂1
P. anxia ♀1
A total of 419 Phyllophaga were taken between 5/1/00 and 7/10/00.
Table 96. Wisconsin, Verona 1999
Blends indicate the ratio of the methyl esters of L-valine/L-isoleucine
100/0 90/10 80/20 60/40 40/60 20/80 10/90 0/100 BLANK
P. fusca ♂1
P. futilis ♂2 1 9 265
P. rugosa ♂2 4
A total of 287 Phyllophaga were taken between 5/13/99 and 6/28/99.
Table 97. Wisconsin, Verona 2000
Blends indicate the ratio of the methyl esters of L-valine/L-isoleucine
100/0 90/10 80/20 60/40 40/60 20/80 10/90 0/100 BLANK
P. fusca ♂1 1
P. futilis ♂1 1 6 322
P. nitida ♂1
P. rugosa ♂2 31 11
P. futilis ♀2
A total of 380 Phyllophaga were taken between 5/9/00 and 6/23/00.
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Table 98a. Synchronic species captured by the same or nearby sex attractant blends, by state.
State Location Year Phyllophaga species Blend
AL Auburn 1997 gracilis, hirticula 0/100
AL Auburn 1998 anxia, gracilis, hirticula, postrema, ulkei 0/100
AL Marion Junction 1999 davisi, futilis 0/100
AL Marion Junction 2000 gracilis var. angulata, forbesi 0/100
CT Vernon 1999 forsteri, fraterna, fusca v/i blends
CT Vernon 2000 forsteri, fraterna, fusca v/i blends
GA Athens 2000 aemula, quercus v/i blends
GA Tifton 1999 glaberrima, uniformis postrema, ulkei v/i blends 0/100
GA Tifton 2000 glaberrima, uniformis v/i blends
IN West Lafayette 2000 balia, futilis 0/100
KS Greensburg #1 2000 corrosa, crassissima, rubiginosa v/i blends
KS Greensburg #2 2001 congrua, crassissima, praetermissa, rubiginosa v/i blends
KS Manhattan #1 2000 crassissima, fusca, rubiginosa, sylvatica, bipartita,futilis v/i blends 0/100
KS Manhattan #2 2001 See Figure 131
KY Lexington 1999 futilis, hirticula 0/100
KY Lexington 2000 See Figure 132 0/100
MA South Amherst 1999 See Figure 133
MA South Amherst 2000 forsteri, fraterna, fusca anxia, longispina v/i blends 0/100
MA Carver 1998 anxia, forsteri v/i blends
MA Carver 2000 anxia, forsteri v/i blends
MA Lakeville 1999 anxia, forsteri, fusca v/i blends
MA Lakeville 2000 anxia, forsteri v/i blends
MA Plympton #1 1996 anxia, forsteri, fraterna v/i blends
MA Plympton #2 1996 anxia, forsteri, fusca v/i blends
ME Lincolnville Center 1999 fraterna, fusca v/i blends
Table 98b. Synchronic species captured by the same or nearby sex attractant blends, by state.
State Location Year Phyllophaga species Blend
ME Lincolnville Center 2000 fraterna, fusca v/i blends
MS Leroy Percy State Park 2000 congrua, hirtiventris v/i blends
MS Sharkey County 2000 congrua, hirtiventris v/i blends
MS Stoneville 2000 congrua, crassissima, hirtiventris v/i blends
NC Raleigh #2 2000 forsteri, micans ephilida, soror v/i blends 0/100
NE Lincoln 1999 crassissima, fusca v/i blends
NE Lincoln 2000 crassissima, fusca v/i blends
NH Madbury 1999 forsteri, fraterna, fusca v/i blends
NH Madbury 2000 forsteri, fraterna, fusca anxia, longispina v/i blends 0/100
NJ Chatsworth #1 1996 anxia,postrema 0/100
NJ Chatsworth #1 1998 anxia,postrema 0/100
NJ Chatsworth #2 2000 anxia,postrema 0/100
NY Bellona 1998 futilis, hirticula 0/100
NY Warwick 2000 balia, drakei 0/100
NY Waterloo #1 1998 anxia, futilis 0/100
NY Waterloo #1 1999 anxia, futilis 0/100
NY Waterloo #1 2000 anxia, futilis 0/100
NY Waterloo #2 1998 anxia, futilis 0/100
PA State College 1999 fraterna, fusca v/i blends
PA State College 2000 fraterna, fusca v/i blends
RI Kingston 1999 anxia, forsteri v/i blends
TX Dallas #1 1998 congrua, crassissima v/i blends
TX Dallas #2 1998 congrua, crassissima v/i blends
TX Dallas #3 1998 congrua, crassissima v/i blends
TX Dallas #4 1998 congrua, crassissima v/i blends
VT Burlington 1999 fraterna, fusca v/i blends
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