THE MICHIGAN ENTOMOLOGICAL SOCIETY ...2007 THE GREAT LAKES ENTOMOLOGIST 103 1Lepidoptera Alert a.k.a. Lepalert, 2806 Linwood Avenue, Royal Oak, MI 48073-3023. (e-mail: [email protected])

THE MICHIGAN ENTOMOLOGICAL SOCIETY2006-2007 OFFICERS

President William Westrate President Elect Stephen Ross Immediate Past President John Douglass Treasurer Martin Andree Secretary Robert Kriegel Member-at-Large (2005-09) Holly Petrillo Member-at-Large (2005-08) Mo Nielson Member-at-Large (2004-07) John Keeler Journal Editor Therese Poland Associate Journal Editor Ronald Priest Newsletter Editor Robert Haack Webmaster Mark O’Brien

The Michigan Entomological Society traces its origins to the old Detroit Entomological Society and was organized on 4 November 1954 to “. . .promote the science of entomology in all its branches and by all feasible means, and to advance cooperation and good fellowship among persons interested in entomology.” The Society attempts to facilitate the exchange of ideas and information in both amateur and professional circles, and encourages the study of insects by youth. Membership in the Society, which serves the North Central States and adjacent Canada, is open to all persons interested in entomology. There are four paying classes of membership for 2008:

Active—annual dues $25.00 Student (to 12th grade)—annual dues $12.00Institutional—annual dues $45.00 Sustaining—annual contribution $35.00 or more Life—$625.00

Dues are paid on a calendar year basis (Jan. 1-Dec. 31).

Memberships accepted before July 1 shall begin on the preceding January 1; memberships accepted at a later date shall begin the following January 1 unless the earlier date is requested and the required dues are paid. All members in good standing receive the Newsletter of the Society and The Great Lakes Entomologist, a semi-annual journal. All active and sustaining members may vote in Society affairs.

All dues and contributions to the Society are deductible for Federal income tax purposes.

SUBSCRIPTION INFORMATION

The journal is published semi-annually as one volume (4 issues) per year. Subscriptions are accepted only on a volume basis. Single copies of The Great Lakes Entomologist are available at $6.00 each, with a 20 percent discount for 25 or more copies sent to a single address.

MICROFILM EDITION: Positive microfilm copies of the current volume of The Great Lakes Ento-mologist will be available at nominal cost, to members and bona fide subscribers of the paper edition only, at the end of each volume year. Please address all orders and inquiries to University Microfilms, Inc., 300 Zeeb Road, Ann Arbor, Michigan 48106, USA.

Inquiries about back numbers, subscriptions and Society business should be directed to the Secretary, Michigan Entomological Society, Department of Entomology, Michigan State University, East Lansing, Michigan 48824-1115, USA. Manuscripts and related correspondence should be directed to the Editor (see inside back cover).

Copyright © 2008. The Michigan Entomological Society.

INSTRUCTIONS FOR AUTHORSSUBJECTS

Papers dealing with any aspect of entomology will be considered for publication in The Great Lakes Entomolo-gist. Appropriate subjects are those of interest to professional and amateur entomologists in the North Central States and Canada, as well as general papers and revisions directed to a larger audience while retaining an interest to readers in our geographic area. All manuscripts are refereed by two reviewers, except for short notes, which are reviewed at the discretion of the Editor.

REQUIREMENTSManuscripts must be typed with line numbers, double-spaced, with 1” margins on 8 1/2 x 11” or equivalent size paper, and submitted in triplicate or emailed to the Editor as an attached file. Please use italics rather than underline. Use subheadings sparingly and set them into paragraphs in boldface. Footnotes (except for authors’ addresses, which must be on the title page, and treated as a footnote), legends, and captions of illustra-tions should be typed on separate sheets of paper. Titles should be concise, identifying the order and family discussed. The author of insect species must be given fully at least once in the abstract and text, but not in the title. If a common name is used for a species or group, it should be in accordance with the common names published by the Entomological Society of America. The format for references must follow that described in the style guidelines used by the Entomological Society of America.

FIGURES & TABLESRemember that the printed page area for The Great Lakes Entomologist is 4.5 x 7 inches. Scale your illustrations accordingly. Photographs should be high resolution digital files (300 dpi) or glossy finish prints (transparencies are not acceptable). Drawings, charts, graphs, and maps must be scaled to permit proper reduction without loss of detail. Figures should also be sent as electronic files and must meet the following criteria: (1) gray-scale im-ages must be submitted as 300 dpi TIFF or EPS files ONLY; (2) line art or graphs must be sent as 600 dpi TIFF or EPS files ONLY. Scanned images should be saved in the native application. Never embed the images in a word-processing document. Captions for figures should be numbered consecutively and typed in order at the end of the manuscript. Captions should not be attached to illustrations or written on the back of images.Tables should be kept as uncluttered as possible, and should fit normally across a page when typeset by the printers. Tables cannot be submitted as Excel files or graphics, but only as text.Contributors should follow the Council of Biology Editors Style Manual, 5th ed., and examine recent issues of The Great Lakes Entomologist for proper format of manuscripts.

MANUSCRIPTS ON DISKManuscripts must be submitted as electronic files along with one printed copy, after they have been accepted for publication. The files may be formatted in any popularly used word-processing program, but must be submitted in Rich Text Format (RTF) or Microsoft Word (DOC) to avoid any translation problems. Special formatting notes for submitting manuscripts on disk: The organizational format for a manuscript is as seen in the recent issues of the journal: TITLE, Author(s), Abstract, Introduction, Methods & Materials, Results, Discussion, Acknowledgments, Literature Cited, Tables, and a List of Figures. Do not use extra spaces between paragraphs or references in the Lit. Cited. The columns of text in tables should be aligned with TABS, not spaces. Some symbols may not translate properly from one computer system to another. Do not use extbols. So long as these symbols are clearly seen in the manuscript, adjustments can be made in the copy sent to the printers.

PAGE CHARGESPapers published in The Great Lakes Entomologist are subject to a page charge of $42.00 per published page. Members of the Society, who are authors without funds from grants, institutions, or industry, and are unable to pay costs from personal funds, may apply to the Society for financial assistance. Application for subsidy must be made at the time a manuscript is initially submitted for publication. Authors will receive a page proof, together with a page charge form. Reprints will be provided as PDF files. Extensive changes to the proof by the author will be billed at a rate of $1.00 per line.

COVER ARTWORKCover art or photographs are desired for upcoming issues. They are published free of charge. We only require that they be suitably prepared as described for images above, and that the subject be identified as accurately as possible.

EDITOR’S ADDRESSAll manuscripts for The Great Lakes Entomologist should be sent to the Editor, Therese Poland, USDA Forest Service, 1407 S. Harrison Rd., Rm. 220, E. Lansing, MI 48823. (email: [email protected]).

OTHER BUSINESSOther correspondence should be directed to the Secretary, Michigan Entomological Society, c/o Dept. of Entomology, Michigan State University, East Lansing, MI 48824-1115.

Vol. 40, Nos. 3 & 4 Fall/Winter 2007

THEGREAT LAKES

ENTOMOLOGIST

The Great Lakes EntomologistPublished by the Michigan Entomological Society

Volume 40 Nos. 3 & 4ISSN 0090-0222

Table of Contents

Using degree-day methodology to ascertain early flight periods of Michigan butterflies and skippers Owen A. Perkins ............................................................................................................. 103

Great diversity of insect floral associates may partially explain ecological success of poison ivy (Toxicodendron radicans subsp. Negundo [Greene] Gillis, Anacardiaceae) David S. Senchina and Keith S. Summerville ..................................................................120

Efficacy of morphological characters for distinguishing nymphs of Epitheca cynosura andEpitheca spinigera (Odonata: Corduliidae) in Wisconsin Robert B. DuBois, Kenneth J. Tennessen, and Matthew S. Berg......................................... 129

Why are there so few insect predators of nuts of American beech (Fagus grandifolia)?Charles E. Williams .........................................................................................................140

Effects of pitfall trap preservation on collections of carabid beetles (Coleoptera: Carabidae) Kenneth W. McCravy and Jason E. Willand ...................................................................... 154

Parasitism of Urophora affinis (Diptera: Tephritidae) by Aprostocetus sp. (Hymenoptera: Eulophidae) in Michigan Jordan M. Marshall .................................................................................................... 166

Arthropods utilizing sticky inflorescences of Cirisium discolor and Penstemon digitalisPatricia A. Thomas .......................................................................................................... 169

An annotated checklist of Wisconsin handsome fungus beetles (Coleoptera: Endomychidae) Michele B. Price and Daniel K. Young ...............................................................................177

A gynandromorph of the March fly, Bibio fraternus (Diptera: Bibionidae)Stephen W. Taber ................................................................................................. NOTE 189

New reports of exotic and native ambrosia and bark beetle species (Coleoptera: Curculionidae: Scolytinae) from Ohio Danielle M. Lightle, Kamal J.K. Gandhi, Anthony I. Cognato, Bryson J. Mosley, David G. Nielsen, and Daniel A. Herms .......................................................... NOTE 194

Designation of a neotype for Mitchell’s satyr, Neonympha mitchellii (Lepidoptera: Nymphalidae)Christopher A. Hamm ........................................................................................... NOTE 201

New record of Vanessa virginiensis (Drury) (Lepidoptera: Nymphalidae) as a host of Thrateles procax (Cresson) (Hymenoptera: Ichneumonidae) David B. MacLean and John Luhman .................................................................. NOTE 203

First occurence of Hippodamia Variegata (Goeze) (Coleoptera: Coccinellidae) in OhioDaniel M. Pavuk, Alan Sundermeier, Sadira Stelzer, Andrea M. Wadsworth, Danielle M. Keeler, Melanie L. Bergolc, and Laura Hughes-Williams ...................... NOTE 205

Cover photo Oarsima poweshiek, (Parker, 1870) (emended), 8 July 2007,

Big Valley Plant Preserve, Buckhorn fen, Rose Township, Oakland County MI.Photo by Dwayne R. Badgero

PUBLISHED BY

THE MICHIGAN ENTOMOLOGICAL

SOCIETY

TH

E G

RE

AT

LAK

ES E

NT

OM

OLO

GIST

Vol. 40, N

os. 3 & 4

Fall/Winter 2007

2007 THE GREAT LAKES ENTOMOLOGIST 103

1Lepidoptera Alert a.k.a. Lepalert, 2806 Linwood Avenue, Royal Oak, MI 48073-3023. (e-mail: [email protected]).

USING DEGREE-DAY METHODOLOGY TO ASCERTAIN EARLY FLIGHT PERIODS OF MICHIGAN BUTTERFLIES AND SKIPPERS

Owen A. Perkins1

ABSTRACTButterflies and skippers have been collected in Michigan for over 130 years

and the accompanying data labels continue to provide significant information. Both collection date and site information for voucher specimens provide the data that is used to ascertain the daily maximum-minimum temperatures for the year in which the specimens were collected. This information may then be used to calculate the degree-day accumulations above a base value over those specific dates.

The Michigan Entomological Society - Michigan Lepidoptera Survey (MLS) is a team of lepidopterists who have endeavored to create a composite database of all voucher specimens in museum and private collections, published data, and submitted data, as well as from surveys conducted throughout the state. This composite data set was used to formulate a first generation of early flight periods of Michigan butterflies and skippers. Accumulated degree-days for predicting the emergence/dispersal and thus the first early flight period are presented for Michigan butterfly and skipper species and subspecies.

____________________

Elements that led to the accumulation of data used to ascertain the grow-ing degree-day values (commonly abbreviated as: degree-day, degday and DD) for each of the Michigan butterfly and skipper populations illustrate the value provided by avocation lepidopterists to the field of entomology. Degree-day is a measure of accumulated heat that can be assigned to each day. Daily values are added together to give an estimate of the amount of development plants, and in this case Lepidoptera, have achieved. The data used in this paper were obtained primarily from the information avocation collectors published or re-corded on individual insect labels or in field notes.

Many collectors have traveled from the Lower Peninsula of Michigan to, and across, the Upper Peninsula to collect butterflies, skippers, and moths. They did this only to find the species had not yet emerged or the flight period, sometimes as short as only a few days, had passed for the season and thus for the year.

About 30 years ago it was discovered by Harry D. King (MLS, personal communication) that he could use a correlation between emergence of the adult stage in Michigan from whatever the over-wintering stage might be (egg, larva, pupa, or adult) and the degree-day accumulation from 1 March of that year. He first used this method for the bog-obligate tundra butterfly, Freija Fritillary, Boloria freija freija, (Thunberg, 1791) (Nymphalidae). Common names used in this paper, though not necessarily approved by the Entomological Society of America, are commonly used by lepidopterists.

The project to determine the early flight period emergence/dispersal date and degree-day value for all of the butterfly and skipper species of Michigan was initiated on 12 July 1997 with the capture of a Poweshiek Skipperling, Oarisma poweshiek (Parker, 1870) (Hesperiidae) (Fig. 1). The degree-day information was used to predict subsequent early flight presence of this species in succeeding years.

104 THE GREAT LAKES ENTOMOLOGIST Vol. 40, Nos. 3 & 4

It is difficult to predict plant growth based on calendar date because tem-perature controls development rate and can vary greatly from month to month and year to year. Instead, degree-days, that are based on actual temperatures, are a simple and accurate way to predict when a certain plant stage will occur. This approach also applies to Lepidoptera. Warmer-than-normal days advance insects’ growth more rapidly, while cooler-than-normal days slows growth com-pared to average. The resulting “thermal time” more consistently predicts when a certain insect stage will occur. One degree-day is one day when the average daily temperature is one degree above the lower developmental threshold (the temperature below which development stops).

Each developmental stage of an organism has its own total heat require-ment. Development can be estimated by accumulating degree-days between the daily high and low temperature thresholds throughout the season. The date to begin accumulating degree-days, known as the biofix date, varies with the species. The biofix date used in this examination has usually been 1 March as it is this time of year that the 50° F (10° C) threshold becomes more constant. Tracking degree-days becomes more vital as the accumulation approaches the degree-day base 50° F value (hereafter referred to as DD50) required for the emergence of the specific species. The DD50 value is used because Maize is a facultative long-night plant and flowers in a certain number of growing degree-days > 50° F in the environment to which it is adapted. The MSU Agricultural Weather Office publishes the values for Maize in terms of DD50 (MSU AWO 2008). The use of the DD50 values has been shown to be effective in arriving at the DD50 value for the emergence of Lepidoptera in the adult stage.

Figure 1. Resident skipper: Poweshiek Skipperling, Oarsima poweshiek (Parker, 1870), 8 July 2007, MNA Big Valley Plant Preserve, Buckhorn fen, Rose Township, Oakland County, MI (photo courtesy of Dwayne R. Badgero).


MATERIALS AND METHODSA database of dates of earliest collection for each species was developed using

data from a collection of voucher specimens, research of collections in the public and private domain, photographic evidence, and capture and release in the field.

Raw data for the degree-day calculation for each species consisted of the earliest dates recorded for each species in the MLS secondary database kept by the author. There were more than 21,000 unique observations for all Michigan lepidopteran species. Earlier dates have been published for a few species in Nielsen (1999) but the years for these were not provided so the earliest dates that included the year were used.

Daily DD50 values for subject dates were obtained from the MSU AWO website (MSU AWO 2008). When the DD50 values were not available from the MSU AWO source, daily maximum and minimum air temperatures from the NOAA (2005) weather station nearest to each location were used to calculate degree-day accumulations when available. When NOAA data were not available, the data were obtained by an alternative source, such as personal records.

Degree-days were calculated above a base of 50° F (DD50) for each obser-vation using the Baskerville-Emin sine method (Baskerville and Emin 1969, Andresen and Harman 1994, Jeffrey A. Andresen, Department of Geography, Michigan State University, personal communication).

Temperature data for the analysis were also obtained from maximum-minimum daily temperatures by recording maximum-minimum daily degrees using a HOBO recording instrument (The HOBO Data Logger Company, Pocas-set, MA). The HOBO Temp data logger is a small, portable, reusable single channel temperature recorder that continuously measures temperature and can be set out in remote locations. The validity of using nearby stations for the actual values at the collecting site was tested in 2003. A HOBO recorder was placed in the field 17 April in Luce County at the west bog field site west of Luce County Road 421 where Boloria frigga saga (Staudinger) (Nymphalidae) and Boloria eunomia dawsoni (W. Barnes & McDunnough) are always found in good number at the peak of their flight season. There was still snow on the road and in the bog and the temperatures in the Upper Peninsula had not yet reached 50° F. The HOBO was retrieved after favorable peak collecting for B. f. saga on 8 June 2003 and indicated a DD50 value of 248. Similarly the MSU Newberry station showed the DD50 as 248 in 2004 for 8 June 2003 indicating that in this particular case using the MSU Newberry station data was a reliable method. The recorded DD50 reported by MSU for 8 June 2003 in 2003 for the Newberry station, however, was given as 233.

The discrepancy in the value reported in different years likely arose be-cause although stations are supposed to take a reading every day and send it into MSU, occasionally data are not reported and a value based on the average of 30 years data is substituted. At the end of the year a hard copy is sent to MSU Agricultural Weather Office and adjustments are made to show the actual readings which generally differ by 10-15° F from the estimated value (King, personal communication). Warm years can be deceptive because degree-days accumulate faster than the insect can respond with faster development (i.e., growth rate is already maximized) thus it results in the appearance of a flight time that is delayed and longer than normal in terms of degree-days.

RESULTS AND DISCUSSIONThe results of accumulating the earliest flight period date and the cor-

responding DD50 values for that date and site for each butterfly and skipper species are provided in Table 1.

106 THE GREAT LAKES ENTOMOLOGIST Vol. 40, Nos. 3 & 4Ta

ble

1. A

ccum

ulat

ed D

egre

e-da

ys a

bove

50°

F fo

r th

e ea

rly

light

per

iod

emer

genc

e/di

sper

sal o

f Mic

higa

n bu

tter

fly a

nd s

kipp

er s

peci

es a

nd

subs

peci

es.

Scie

ntifi

c na

mea

Col

lect

or

MLS

Ear

ly D

ateb

Cou

nty

DD

50F

Flig

htc

HES

PERI

IDAE

Epa

rgyr

eus

clar

us c

laru

s (C

ram

er)

Kuh

lman

, Rog

er

2 M

ay 1

999

Was

hten

aw

170

RU

rban

us p

rote

us (L

inna

eus)

N

ewco

mb,

Will

iam

W.

Prio

r 19

13

Way

ne

N/A

S

Acha

laru

s ly

ciad

es (G

eyer

) K

uhlm

an, R

oger

9

May

200

1 W

asht

enaw

28

6 R

Thor

ybes

bat

hyllu

s (J

. E. S

mith

) M

oore

, She

rman

5

May

194

1 Li

ving

ston

19

3 R

Thor

ybes

pyl

ades

pyl

ades

(Scu

dder

) Pe

rkin

s, O

wen

A.

12

May

200

0 *

Was

hten

aw

320

RPh

olio

sora

cat

ullu

s (F

abri

cius

) Pe

rkin

s, O

wen

A.

5 M

ay 2

000

* Ca

ss

202

RE

rynn

is ic

elus

(Scu

dder

& B

urge

ss)

Perk

ins,

Ow

en A

. 24

Apr

199

8 O

akla

nd

157

RE

rynn

is b

rizo

bri

zo (B

oisd

uval

& L

e Co

nte)

K

uhlm

an, R

oger

18

Apr

200

0 W

asht

enaw

96

R

Ery

nnis

juve

nalis

juve

nalis

(Fab

rici

us)

Perk

ins,

Ow

en A

. 20

Apr

199

8 O

akla

nd

140

RE

rynn

is h

orat

ius

(Scu

dder

& B

urge

ss)

Kuh

lman

, Rog

er

29 A

pr 2

000

Was

hten

aw

140

RE

rynn

is m

artia

lis (S

cudd

er)

Perk

ins,

Ow

en A

. 13

May

195

1 Ca

lhou

n 17

8 R

Ery

nnis

fune

ralis

(Scu

dder

& B

urge

ss)

Gra

nlun

d, J

ames

L.

18 A

ug 2

001

Chip

pew

a 14

54

SE

rynn

is b

aptis

iae

(W. F

orbe

s)

Bial

ecki

, Mar

tin J

. 22

Apr

200

6 M

onro

e 15

2 R

Ery

nnis

luci

lius

(Scu

dder

& B

urge

ss)

Perk

ins,

Ow

en A

. 5

May

200

0 M

onro

e 19

7 R

Ery

nnis

per

sius

per

sius

(Scu

dder

) (1st

bro

od)

Oos

ting

Dan

iel S

. 1

May

197

6 Al

lega

n 16

9 R

Ery

nnis

per

sius

per

sius

(Scu

dder

) (2nd

bro

od)

Perk

ins,

Ow

en A

. 8

Jul 2

007

Oak

land

11

89

RPy

rgus

cen

taur

eae

wya

ndot

(W. H

. Edw

ards

) N

iels

en, M

ogen

s C.

3

May

198

1 M

ontc

alm

15

1 R

Pyrg

us c

omm

unis

com

mun

is (G

rote

) Fa

rmer

, Joh

n C.

9

Apr 2

004

Hill

sdal

e 55

R

Car

tero

ceph

alus

pal

aem

on m

anda

n (W

. H. E

dwar

ds)

Perk

ins,

Ow

en A

. 15

May

199

8 Sc

hool

craf

t 13

5 R

Ancy

loxy

pha

num

itor

(Fab

rici

us)

Farm

er, J

ohn

C.

9 Ap

r 200

4 H

illsd

ale

55

RO

aris

ma

pow

eshe

ik (P

arke

r)

New

com

b, W

illia

m W

. 9

Jun

1930

K

ent

789

RO

aris

ma

pow

eshi

ek (P

arke

r) e

men

ded

Perk

ins,

Ow

en A

. 22

Jun

199

8 O

akla

nd

849

RTh

ymel

icus

line

ola

lineo

la (O

chse

nhei

mer

) Pe

rkin

s, O

wen

A.

1 Ju

n 19

98 *

O

akla

nd

605

RC

alpo

des

ethl

ius

(Sto

ll)

Wea

ver,

Scot

t 29

May

199

9 Ja

ckso

n 47

5 S

Pano

quin

a oc

ola

ocol

a (W

. H. E

dwar

ds)

Kuh

lman

, Rog

er

2 O

ct 2

005

Mon

roe

2985

S

Ambl

ysci

rtes

heg

on (S

cudd

er)

Kuh

lman

, Rog

er

24 M

ay 2

007

* Li

ving

ston

39

0 R

Ambl

ysci

rtes

via

lis (W

. H. E

dwar

ds)

Her

ig, T

erry

L.

5 M

ay 2

000

Cass

20

2 R

Nas

tra

lher

min

ier

(Lat

reill

e)

Koe

hn, L

arry

C.

17 J

ul 1

983

Cass

13

73

SLe

rode

s eu

fala

euf

ala

(W. H

. Edw

ards

) D

reis

bach

, Rob

ert R

. 25

Aug

195

9 O

nton

agon

14

08

S

2007 THE GREAT LAKES ENTOMOLOGIST 107Ta

ble

1. C

ontin

ued.

Scie

ntifi

c na

mea

Col

lect

or

MLS

Ear

ly D

ateb

Cou

nty

DD

50F

Flig

htc

Lere

ma

acci

us (J

. E. S

mith

) H

erig

, Ter

ry L

. 4

Sep

1998

Le

naw

ee

2656

S

Hyl

ephi

la p

hyle

us p

hyle

us (D

rury

) K

uhlm

an, R

oger

14

Jun

199

9 W

asht

enaw

82

6 M

Hes

peri

a co

mm

a la

uren

tina

(Lym

an)

John

son,

Kyl

e E.

27

Jul

200

3 *

Del

ta

1065

R

Hes

peri

a ot

toe

(W. H

. Edw

ards

) Ba

logh

, Geo

rge

J.

1 Ju

n 19

87

Alle

gan

663

RH

espe

ria

leon

ardu

s le

onar

dus

(T. H

arri

s)

New

com

b, W

illia

m W

. 3

Jul 1

930

Dic

kins

on

742

RH

espe

ria

met

ea m

etea

Scu

dder

Pe

rkin

s, O

wen

A.

5 M

ay 2

000

Alle

gan

202

RH

espe

ria

sass

acus

sas

sacu

s (T

. Har

ris)

Pe

rkin

s, O

wen

A.

29 M

ay 1

999

* Ar

enac

35

7 R

Polit

es p

ecki

us p

ecki

us (W

. Kir

by)

Hol

zman

, Ric

hard

W.

7 Ap

r 195

8 Li

ving

ston

21

R

Polit

es th

emis

tocl

es th

emis

tocl

es (L

atre

ille)

W

arcz

ynsk

i, Vi

rgil

J.

11 M

ay 1

963

Lape

er

201

RPo

lites

ori

gene

s or

igen

es (F

abri

cius

) K

uhlm

an, R

oger

16

Jun

200

2 *

Was

hten

aw

633

RPo

lites

mys

tic m

ystic

(W. H

. Edw

ards

) W

arcz

ynsk

i, Vi

rgil

J.

5 M

ay 1

962

Aren

ac

131

RW

alle

ngre

nia

eger

emet

(Scu

dder

) M

oore

, She

rman

8

Jun

1939

Li

ving

ston

54

0 R

Pom

peiu

s ve

rna

(W. H

. Edw

ards

) Re

inoe

hl, J

ack

4 Ju

n 20

04

Gra

tiot

519

RAt

alop

edes

cam

pest

ris

huro

n (W

. H. E

dwar

ds)

unkn

own

15 S

ep 1

999

* W

asht

enaw

26

50

SPo

anes

hob

omok

hob

omok

(T. H

arri

s)

Kuh

lman

, Rog

er

12 M

ay 2

000

Was

hten

aw

320

RPo

anes

zab

ulon

(Boi

sduv

al &

Le

Cont

e)

Nie

lsen

, Mog

ens

C.

28 M

ay 1

989

St. J

osep

h 50

2 R

Poan

es m

assa

soit

mas

saso

it (S

cudd

er)

Moo

dy, P

hilip

E.

29 M

ay 1

938

Oak

land

34

4 R

Poan

es v

iato

r vi

ator

(W. H

. Edw

ards

) Re

inoe

hl, J

ack

28 J

un 2

007

Gra

tiot

1133

R

Anat

ryto

ne lo

gan

loga

n (W

. H. E

dwar

ds)

Kuh

lman

, Rog

er

17 J

un 2

001

* W

asht

enaw

76

5 R

Eup

hyes

con

spic

ua c

onsp

icua

(W. H

. Edw

ards

) W

arcz

ynsk

i, Vi

rgil

J.

6 Ju

n 19

62

Lape

er

610

RE

uphy

es d

ion

(W. H

. Edw

ards

) W

arcz

ynsk

i, Vi

rgil

J.

17 J

un 1

963

Sagi

naw

77

7 R

Eup

hyes

duk

esi d

ukes

i (Li

ndse

y)

Kle

itch

(Fet

tinge

r), J

enni

fer

5 Ju

n 20

03

Mus

kego

n 50

1 R

Eup

hyes

bim

acul

a bi

mac

ula

(Gro

te &

Rob

inso

n)

Wag

ner,

War

ren

H. J

r. 4

Jun

1956

W

asht

enaw

31

1 R

Eup

hyes

ves

tris

met

acom

et (T

. Har

ris)

Le

ski,

Mic

hael

L.

29 M

ay 2

006

Ont

onag

on

302

RAt

ryto

nops

is h

iann

a hi

anna

(Scu

dder

) H

odge

s, R

onal

d W

. 21

May

195

8 Cr

awfo

rd

183

R

PAPI

LIO

NID

AEB

attu

s ph

ileno

r ph

ileno

r (L

inna

eus)

(1st

broo

d)

War

czyn

ski,

Virg

il J.

20

Apr

196

6 Le

naw

ee

155

RB

attu

s ph

ileno

r ph

ileno

r (L

inna

eus)

(2nd

broo

d)

Chur

chill

, Mar

k 6

Jul 2

008

Alle

gan

926

RE

uryt

ides

mar

cellu

s (C

ram

er) (

1st b

rood

) N

iels

en, M

ogen

s C.

28

Apr

195

5 Be

rrie

n 17

5 R


ble

1. C

ontin

ued.

Scie

ntifi

c na

mea

Col

lect

or

MLS

Ear

ly D

ateb

Cou

nty

DD

50F

Flig

htc

Eur

ytid

es m

arce

llus

(Cra

mer

) (2nd

bro

od)

Hub

bell,

The

odor

e H

. 26

Jun

191

9 Be

rrie

n 89

8 R

Papi

lio m

acha

on h

udso

nian

us A

. Cla

rk

Boga

r, D

anie

l S.

9 Ju

n 19

91

Chip

pew

a 53

3 S

Papi

lio p

olyx

enes

ast

eriu

s (S

toll)

(1st

broo

d)

Star

key,

Don

ald

J.

14 A

pr 1

997

* M

acom

b 54

R

Papi

lio p

olyx

enes

ast

eriu

s (S

toll)

(2nd

broo

d)

Perk

ins,

Ow

en A

. 16

Jun

195

1 O

akla

nd

630

RPa

pilio

cre

spho

ntes

Cra

mer

(1st b

rood

) Be

ss, J

ames

A.

25 A

pr 1

983

Jack

son

168

RPa

pilio

cre

spho

ntes

Cra

mer

(2nd

bro

od)

Perk

ins,

Ow

en A

. 22

Jun

199

7 O

akla

nd

637

RPa

pilio

can

aden

sis

Roth

schi

ld &

Jor

dan

Nie

lsen

, Mog

ens

C.

28 A

pr 2

000

Ots

ego

83

RPa

pilio

gla

ucus

gla

ucus

Lin

naeu

s Bi

alec

ki, M

artin

J.

15 A

pr 2

000

Was

hten

aw

84

RPa

pilio

troi

lus

troi

lus

Linn

aeus

(1st

broo

d)

unkn

own

22 A

pr 1

914

Ingh

am

134

RPa

pilio

troi

lus

troi

lus

Linn

aeus

(2nd

broo

d)

Legg

e, J

ohn

T.

12 J

ul 1

994

Kal

amaz

oo

1255

R

PIER

IDAE

Nat

halis

iole

Boi

sduv

al

Ross

, Ste

phen

18

Jul

200

1 G

ogeb

ic

1068

M

/RE

urem

a m

exic

ana

mex

ican

a (B

oisd

uval

) G

roco

ff, M

. 7

Jun

1977

Li

ving

ston

87

4 S

Pyri

sitia

lisa

lisa

(Boi

sduv

al &

Le

Cont

e)

unkn

own

15 M

ay 1

964

Ingh

am

200

M/R

Abae

is n

icip

pe (C

ram

er)

Dee

ring

, Mar

k 20

Jun

199

6 K

alam

azoo

75

0 S

Col

ias

philo

dice

phi

lodi

ce G

odar

t Ya

tes,

Art

hur A

. 4

Apr 1

943

Pres

que

Isle

10

1 R

Col

ias

eury

them

e (B

oisd

uval

) Bi

alec

ki, M

artin

J.

22 A

pr 2

006

Mon

roe

152

RC

olia

s in

teri

or S

cudd

er

Perk

ins,

Ow

en A

. 20

Jun

199

8 *

Ots

ego

653

RZe

rene

ces

onia

ces

onia

(Sto

ll)

New

man

, Joh

n W

. 23

May

193

9 O

akla

nd

239

SPh

oebi

s se

nnae

sen

nae

= eu

bule

(Lin

naeu

s)

Farm

er, J

ohn

C.

21 M

ay 1

999

Was

hten

aw

390

M/R

Phoe

bis

phile

a ph

ilea

(Lin

naeu

s)

Beeb

e, R

alph

25

Aug

195

1 W

ayne

24

12

SE

uchl

oe a

uson

ides

may

i F. C

herm

ock

& R

. Che

rmoc

k N

iels

en, M

ogen

s C.

11

May

198

7 K

ewee

naw

46

R

Euc

hloe

oly

mpi

a =

rosa

(W. H

. Edw

ards

) N

iels

en, M

ogen

s C.

23

Apr

195

5 Be

rrie

n 15

2 R

Pier

is o

lera

cea

oler

acea

(T. H

arri

s) (1

st br

ood)

Kuh

lman

, Rog

er

11 A

pr 1

998

Was

hten

aw

107

RPi

eris

ole

race

a ol

erac

ea (T

. Har

ris)

(2nd

broo

d)

M

cAlp

ine,

Wilb

ur S

. 21

Jun

193

6 Ca

ss

700

RPi

eris

vir

gini

ensi

s vi

rgin

iens

is W

. H. E

dwar

ds

N

iels

en, M

ogen

s C.

24

Apr

199

0 Ca

ss

145

RPi

eris

rap

ae r

apae

(Lin

naeu

s)

Kuh

lman

, Rog

er

23 M

ar 2

000

Was

hten

aw

40

RPo

ntia

pro

todi

ce (B

oisd

uval

& L

e Co

nte)

W

agne

r, W

arre

n H

. Jr.

18 A

pr 1

981

* St

. Jos

eph

133

RPo

ntia

occ

iden

talis

occ

iden

talis

(Rea

kirt

)

Nie

lsen

, Mog

ens

C.

19 J

un 1

989

Mon

tcal

m

917

SAs

cia

mon

uste

phi

leta

(Fab

rici

us)

Mon

nier

, Fra

ncis

X.

N/A

N

/A

N/A

S


ble

1. C

ontin

ued.

Scie

ntifi

c na

mea

Col

lect

or

MLS

Ear

ly D

ateb

Cou

nty

DD

50F

Flig

htc

LYCA

ENID

AEFe

nise

ca ta

rqui

nius

tarq

uini

us (F

abri

cius

) K

uhlm

an, R

oger

26

Apr

200

0 W

asht

enaw

12

4 R

Lyca

ena

phla

eas

hypo

phla

eas

= am

eric

ana

T. H

arri

s K

uhlm

an, R

oger

1

May

199

9 *

Was

hten

aw

162

RLy

caen

a hy

llus

(Cra

mer

) M

cKen

ney,

M. J

. 25

Apr

196

2 Li

ving

ston

68

R

Lyca

ena

epix

anth

e m

ichi

gane

nsis

Raw

son

New

man

, Joh

n W

. 4

Jun

1946

O

akla

nd

407

RLy

caen

a do

rcas

mic

huro

n Sc

ott

Smith

, Tod

d 6

Jul 2

007

Oak

land

11

44

RLy

caen

a do

rcas

dor

cas

W. K

irby

Fi

sche

r, Ro

land

L.

10 J

un 1

959

Barr

y 70

1 R

Lyca

ena

hello

ides

(Boi

sduv

al)

Hod

ges,

Ron

ald

W.

14 M

ay 1

955

Mon

tcal

m

300

RSa

tyri

um a

cadi

ca a

cadi

ca (W

. H. E

dwar

ds)

McK

enne

y, M

. J.

25 A

pr 1

962

Livi

ngst

on

68

RSa

tyri

um ti

tus

titus

(Fab

rici

us)

McK

enne

y, M

. J.

25 A

pr 1

962

Livi

ngst

on

68

RSa

tyri

um e

dwar

dsii

edw

ards

ii (G

rote

& R

obin

son)

Nie

lsen

, Mog

ens

C.

10 J

un 2

004

New

aygo

56

3 R

Saty

rium

cal

anus

fala

cer

(God

art)

N

iels

en, M

ogen

s C.

25

Apr

195

2 Li

ving

ston

14

8 R

Saty

rium

car

yaev

orus

(McD

unno

ugh)

M

cKen

ney,

M. J

. 25

Apr

196

2 Li

ving

ston

68

R

Saty

rium

lipa

rops

str

igos

a (T

. Har

ris)

N

ewm

an, J

ohn

H.

3 Ju

n 19

44

Wex

ford

24

7 R

Saty

rium

favo

nius

ont

ario

(W. H

. Edw

ards

) O

ostin

g D

anie

l S.

28 J

un 1

975

Lena

wee

93

0 S

Cal

loph

rys

augu

stin

us a

ugus

tinus

(Wes

twoo

d)

Kuh

lman

, Rog

er

14 A

pr 2

003

Was

hten

aw

88

RC

allo

phry

s po

lios

polio

s (C

ook

& F

. Wat

son)

Pe

rkin

s, O

wen

A.

27 A

pr 2

000

* Ch

ippe

wa

49

RC

allo

phry

s ir

us ir

us (G

odar

t)

Nie

lsen

, Mog

ens

C.

25 A

pr 1

987

Alle

gan

243

RCa

lloph

rys

henr

ici h

enri

ci (G

rote

& R

obin

son)

N

iels

en, M

ogen

s C.

28

Apr

198

1 M

ontc

alm

15

4 R

Cal

loph

rys

niph

on c

lark

i (T.

Fre

eman

) W

arcz

ynsk

i, Vi

rgil

J.

18 A

pr 1

968

Aren

ac

102

RC

allo

phry

s er

ypho

n er

ypho

n (B

oisd

uval

) Pe

rkin

s, O

wen

A.

27 A

pr 2

000

Chip

pew

a 49

R

Cal

ycop

is c

ecro

ps (F

abri

cius

) M

onni

er, F

ranc

is X

. 10

Jul

188

2 Be

rrie

n N

/A

SSt

rym

on m

elin

us h

umul

i (T.

Har

ris)

N

iels

en, M

ogen

s C.

23

Apr

195

5 Be

rrie

n 15

2 R

Parr

hasi

us m

-alb

um (B

oisd

uval

& L

e Co

nte)

Ta

ggar

t, Jo

hn

12 A

ug 1

964

Mus

kego

n 20

17

SE

rora

laet

a (W

. H. E

dwar

ds) (

1st b

rood

) El

sner

, Elw

in A

. 9

May

199

8 Le

elan

au

122

RE

rora

laet

a (W

. H. E

dwar

ds) (

2nd b

rood

) O

ostin

g, D

anie

l S.

21 J

ul 1

981

Emm

et

909

RC

upid

o co

myn

tas

com

ynta

s (G

odar

t)

Bial

ecki

, Mar

tin J

. 22

Apr

200

6 M

onro

e 15

2 R

Cup

ido

amyn

tula

am

yntu

la (B

oisd

uval

) M

oore

, She

rman

3

May

192

0 M

acki

nac

43

M/R

Cel

astr

ina

luci

a lu

cia

(W. K

irby

) Be

ss, J

ames

A.

18 A

pr 1

987

Ots

ego

81

RC

elas

trin

a la

don

(Cra

mer

) K

uhlm

an, R

oger

1

Apr 2

000

Was

hten

aw

63

R


ble

1. C

ontin

ued.

Scie

ntifi

c na

mea

Col

lect

or

MLS

Ear

ly D

ateb

Cou

nty

DD

50F

Flig

htc

Cel

astr

ina

sero

tina

Pavu

lan

& D

. Wri

ght

Her

ig J

r., E

dwar

d 1

May

200

6 Cl

into

n 17

0 R

Cel

astr

ina

negl

ecta

(W. H

. Edw

ards

) St

inso

n, W

alte

r C.

31 M

ay 1

930

Was

hten

aw

400

RG

lauc

opsy

che

lygd

amus

cou

peri

Gro

te

Hod

ges,

Ron

ald

W.

30 A

pr 1

965

Mon

tcal

m

134

RE

chin

argu

s is

ola

(Rea

kirt

) Sw

ales

, Joh

n 29

May

199

6 W

asht

enaw

33

0 M

/RPl

ebej

us id

as n

abok

ovi (

Mas

ters

) N

iels

en, M

ogen

s C.

3

Jun

1987

Al

ger

149

RPl

ebej

us m

elis

sa s

amue

lis (N

abok

ov) (

1st br

ood)

Her

ig J

r., E

dwar

d

5 M

ay 1

989

Alle

gan

213

RPl

ebej

us m

elis

sa s

amue

lis (N

abok

ov) (

2nd br

ood)

Oos

ting,

Dan

iel S

. 22

Jun

198

2 Al

lega

n 68

1 R

Pleb

ejus

mel

issa

sam

uelis

(Nab

okov

) (pe

ak)

Perk

ins,

Ow

en A

. 7

Jul 2

007

Mon

tcal

m

1325

R

Pleb

ejus

sae

piol

us a

mic

a (W

. H. E

dwar

ds)

New

man

, Joh

n W

. 20

May

194

1 O

scod

a 13

4 R

RIO

DIN

IDAE

Cal

ephe

lis m

utic

um M

cAlp

ine

Wag

ner,

War

ren

H. J

r. 21

Jun

195

4 W

asht

enaw

99

2 R

NYM

PHAL

IDAE

Liby

thea

na c

arin

enta

bac

hman

ii K

irtla

nd

Oos

ting,

Dan

iel S

. 1

Jun

1970

O

ttaw

a 39

0 M

Dan

aus

plex

ippu

s pl

exip

pus

(Lin

naeu

s)

Bial

ecki

, Mar

tin J

. 21

Apr

200

6 W

asht

enaw

13

0 M

/RD

anau

s gi

lippu

s be

reni

ce (C

ram

er)

Kuh

lman

, Rog

er

17 J

ul 2

003

Was

hten

aw

1277

S

Lim

eniti

s ar

them

is a

rthe

mis

(Dru

ry)

unkn

own

21 M

ay 1

902

Alge

r N

/A

RLi

men

itis

arth

emis

art

hem

is (D

rury

) W

arcz

ynsk

i, Vi

rgil

J.

14 J

un 1

964

Mar

quet

te

418

RLi

men

itis a

rthe

mis

= p

rose

rpin

a W

. H. E

dwar

ds

Lesk

i, M

icha

el L

. 29

May

200

6 O

nton

agon

30

2 R

Lim

eniti

s ar

them

is =

alb

ofas

ciat

a N

ewco

mb

New

man

, Joh

n H

. 16

Jun

194

4 O

scod

a 34

8 R

Lim

eniti

s a. r

ubro

fasc

iata

(Bar

nes &

McD

unno

ugh)

Pe

rkin

s, O

wen

A.

17 A

ug 1

999

Mac

kina

c 13

00

RLi

men

itis

arth

emis

ast

yana

x (F

abri

cius

) K

uhlm

an, R

oger

19

May

199

8 W

asht

enaw

47

3 R

Lim

eniti

s ar

chip

pus

arch

ippu

s (C

ram

er)

Kuh

lman

, Rog

er

19 M

ay 2

001

Was

hten

aw

397

RAg

raul

is v

anill

ae n

igri

or M

iche

ner

Kuh

lman

, Rog

er

21 J

ul 1

998

Was

hten

aw

1881

S

Eup

toie

ta c

laud

ia (C

ram

er)

Kuh

lman

, Rog

er

26 A

pr 2

004

Was

hten

aw

176

M/R

Bol

oria

euno

mia

daw

soni

(W. B

arne

s & M

cDun

noug

h) P

erki

ns, O

wen

A.

27 M

ay 1

998

Chip

pew

a 32

2 R

Bol

oria

sel

ene

myr

ina

(Cra

mer

) W

arcz

ynsk

i, Vi

rgil

J.

4 M

ay 1

962

Bay

123

RB

olor

ia s

elen

e at

roco

stal

is (H

uard

) Pe

rkin

s, O

wen

A.

15 M

ay 1

998

Scho

olcr

aft

135

RB

olor

ia b

ello

na b

ello

na (F

abri

cius

) K

uhlm

an, R

oger

25

Apr

199

9 W

asht

enaw

12

0 R


ble

1. C

ontin

ued.

Scie

ntifi

c na

mea

Col

lect

or

MLS

Ear

ly D

ateb

Cou

nty

DD

50F

Flig

htc

Bol

oria

frig

ga s

aga

(Sta

udin

ger)

Pe

rkin

s, O

wen

A.

15 M

ay 1

998

Luce

13

5 R

Bol

oria

frei

ja fr

eija

(Thu

nber

g)

Perk

ins,

Ow

en A

. 9

May

199

8 *

Dic

kins

on

81

RSp

eyer

ia c

ybel

e cy

bele

(Fab

rici

us)

Kuh

lman

, Rog

er

20 M

ay 1

998

Was

hten

aw

496

RSp

eyer

ia c

ybel

e kr

autw

urm

i (W

. Hol

land

) U

nkno

wn

25 J

un 1

910

* Al

ger

902

RSp

eyer

ia a

phro

dite

aph

rodi

te (F

abri

cius

) M

artin

at,P

eter

J.

6 Ju

n 19

74

Alle

gan

665

RSp

eyer

ia a

phro

dite

alc

estis

(W. H

. Edw

ards

) Pe

rkin

s, O

wen

A.

3 Ju

l 200

0 Ba

rry

942

RSp

eyer

ia id

alia

idal

ia (D

rury

) K

ing,

Har

ry D

. 25

Jun

197

7 *

St. J

osep

h 11

99

RSp

eyer

ia a

tlant

is a

tlant

is (W

. H. E

dwar

ds)

War

czyn

ski,

Virg

il J.

26

May

196

3 O

tseg

o 21

8 R

Aste

roca

mpa

cel

tis c

eltis

(Boi

sduv

al &

Le

Cont

e)

Kuh

lman

, Rog

er

2 Ju

n 20

07

Was

hten

aw

620

RAs

tero

cam

pa c

lyto

n cl

yton

(Boi

sduv

al &

Le

Cont

e)

Kin

g, H

arry

D.

30 M

ay 1

977

Barr

y 55

0 R

Vane

ssa

virg

inie

nsis

(Dru

ry)

Kuh

lman

, Rog

er

8 Ap

r 200

1 W

asht

enaw

0

RVa

ness

a ca

rdui

(Lin

naeu

s)

Ross

, Ste

phen

18

Apr

200

5 M

ecos

ta

112

M/R

Vane

ssa

atal

anta

rub

ria

(Fru

hsto

rfer

) K

uhlm

an, R

oger

5

Apr 1

998

Was

hten

aw

99

M/R

Agla

is m

ilber

ti m

ilber

ti (G

odar

t)

Kuh

lman

, Rog

er

5 M

ar 2

000

Was

hten

aw

0 R

Nym

phal

is l-

albu

m j-

albu

m (B

oisd

uval

& L

e Co

nte)

K

uhlm

an, R

oger

12

Mar

200

2 W

asht

enaw

0

RN

ymph

alis

cal

iforn

ica

(Boi

sduv

al)

Voss

, Edw

ard

G.

4 Se

p 19

45

Emm

et/M

ason

18

35

SN

ymph

alis

ant

iopa

ant

iopa

(Lin

naeu

s)

Kuh

lman

, Rog

er

28 F

eb 2

004

Was

hten

aw

2 R

Poly

goni

a in

terr

ogat

ioni

s (F

abri

cius

) Pe

rkin

s, O

wen

A.

9 Ju

n 19

69

Oak

land

46

8 R

Poly

goni

a in

terr

ogat

ioni

s =

umbr

osa

(Lin

tner

) K

uhlm

an, R

oger

14

Apr

200

3 W

asht

enaw

88

R

Poly

goni

a co

mm

a co

mm

a (T

. Har

ris)

Pe

rkin

s, O

wen

A.

28 F

eb 1

998

Oak

land

7

RPo

lygo

nia

com

ma

= dr

yas

(W. H

. Edw

ards

) Be

ss, J

ames

A.

19 A

pr 1

980

Jack

son

139

RPo

lygo

nia

saty

rus

neom

arsy

as d

os P

asso

s N

iels

en, M

ogen

s C.

17

May

200

1 *

Iron

21

8 R

Poly

goni

a pr

ogne

(Cra

mer

) Pe

rkin

s, O

wen

A.

7 M

ar 2

000

Oak

land

32

R

Poly

goni

a pr

ogne

= l-

arge

nteu

m S

cudd

er

Don

ahue

, Jul

ian

P.

11 A

pr 1

964

Ingh

am

24

RPo

lygo

nia

grac

ilis

grac

ilis

(Gro

te &

Rob

inso

n)

N

iels

en, M

ogen

s C.

29

May

196

0 Sc

hool

craf

t 17

8 R

Poly

goni

a fa

unus

faun

us (W

. H. E

dwar

ds)

Don

ahue

, Jul

ian

P.

30 M

ar 1

944

Ots

ego

0 R

Juno

nia

coen

ia c

oeni

a (H

ubne

r)

Kuh

lman

, Rog

er

12 M

ay 2

000

Was

hten

aw

320

M/R

Juno

nia

coen

ia =

ros

a (W

hitt

aker

& D

. Sta

lling

s)

Nie

lsen

, Mog

ens

C.

17 S

ep 2

002

Cass

27

00

M/R

Juno

nia

geno

veva

(Cra

mer

) K

nigh

t, K

en

20 O

ct 2

007

Ken

t 31

55

SE

uphy

drya

s ph

aeto

n ph

aeto

n (D

rury

) K

ing,

Har

ry D

. 28

May

199

5 *

Ingh

am

465

R


ble

1. C

ontin

ued.

Scie

ntifi

c na

mea

Col

lect

or

MLS

Ear

ly D

ateb

Cou

nty

DD

50F

Flig

htc

Chl

osyn

e ny

ctei

s ny

ctei

s (E

. Dou

bled

ay)

McK

enne

y, M

. J.

25 A

pr 1

962

Livi

ngst

on

68

RC

hlos

yne

gorg

one

carl

ota

(Rea

kirt

) G

aige

, Fre

deri

ck M

. 26

May

193

4 Io

sco

221

RC

hlos

yne

harr

isii

harr

isii

(Scu

dder

) Pe

rkin

s, O

wen

A.

1 Ju

n 19

98

Oak

land

60

5 R

Phyc

iode

s th

aros

thar

os (D

rury

) M

cKen

ney,

M. J

. 25

Apr

196

2 Li

ving

ston

68

R

Phyc

iode

s co

cyta

sel

enis

(W. K

irby

) Cu

thre

ll, D

avid

L.

30 A

pr 1

995

Kew

eena

w

37

RPh

ycio

des

bate

sii b

ates

ii (R

eaki

rt)

McA

lpin

e, W

ilbur

S.

4 M

ay 1

941

* M

ontm

oren

cy

101

RAn

aea

andr

ia S

cudd

er

Stin

son,

Wal

ter C

. 8

May

193

2 W

asht

enaw

20

6 S

Leth

e an

thed

on a

nthe

don

A. C

lark

St

arke

y, D

onal

d J.

18

Jun

199

8 M

acom

b 80

7 R

Leth

e an

thed

on b

orea

lis A

. Cla

rk

Perk

ins,

Ow

en A

. 22

Jun

200

7 Al

ger

610

RLe

the

creo

la (S

kinn

er)

Bruc

e, D

avid

N

/A

N/A

N

/A

SLe

the

eury

dice

eur

ydic

e (L

inna

eus)

Ex

, Jac

ob C

. 25

May

200

5 O

akla

nd

303

RLe

the

appa

lach

ia le

euw

i Gat

relle

& A

rbog

ast

Star

key,

Don

ald

J.

14 J

un 1

998

Oak

land

77

9 R

Coe

nony

mph

a tu

llia

inor

nata

W. H

. Edw

ards

K

atz,

Ste

ve

28 M

ay 2

007

Oak

land

45

8 R

Neo

nym

pha

mitc

helli

i mitc

helli

i Fre

nch

Har

vey,

Don

ald

J.

14 J

un 1

974

Jack

son

580

RM

egis

to c

ymel

a cy

mel

a (C

ram

er)

Kuh

lman

, Rog

er

16 M

ay 1

998

* W

asht

enaw

40

9 R

Cer

cyon

is p

egal

a al

ope

(Fab

rici

us)

Hun

ter,

Jam

es

1 Ju

n 19

78

Oak

land

51

8 R

Cer

cyon

is p

egal

a ne

phel

e (W

. Kir

by)

Unk

now

n 7

Jun

1940

In

gham

63

9 R

Ere

bia

disc

oida

lis d

isco

idal

is (W

. Kir

by)

Badg

ero,

Dw

ayne

R.

3 M

ay 2

007

Chip

pew

a 76

R

Oen

eis

jutta

asc

erta

Mas

ters

& S

oren

son

Ross

, Ste

phen

20

May

200

6 D

elta

15

8 R

Oen

eis

chry

xus

stri

gulo

sa M

cDun

noug

h K

ing,

Har

ry D

. 2

May

199

9 N

eway

go

143

RO

enei

s m

acou

nii (

W. H

. Edw

ards

) N

iels

en, M

ogen

s C.

16

Jun

198

6 K

ewee

naw

43

1 R

a Ar

rang

emen

t by

Pelh

am e

t al.

(200

8).

b M

LS is

Mic

higa

n Le

pido

pter

a Su

rvey

dat

a.c F

light

key

is: R

=Res

iden

t, M

=Mig

rant

, M/R

=Mig

rant

/Res

iden

t, S=

Stra

y.* Ea

rly

date

dat

a do

es n

ot h

ave

year

pro

vide

d, th

eref

ore

alte

rnat

e ea

rly

date

with

yea

r is

used

.


Although insect phenology modeling using degree days is a common strat-egy for managing agricultural pests, it is rarely used to predict the phenology of threatened or endangered insects. The present study describes an analytical method using historical records that produces a phenological model adequate for conducting field surveys of some insect species (Robert D. Kriegel and Mogens C. Nielsen, Michigan State University, personal communication).

It is important to also ascertain the peak flight period as it may be derived from the data used to determine the early flight period date and documented observations of large explosive emergences. This has been done to some extent for the Federally Endangered Karner Blue Plebejus melissa samuelis (Nabokov, 1944) (Lycaenidae) (Nielsen 1999; Fig. 2).

The degree-day predictions for Karner Blue emergence were tested in 2007. On 7 July 2007 a team of MLS members and others encountered a peak emergence of Karner Blue while surveying in Montcalm County at a site where the larval host plant, lupine, Lupinus perennis L., was previously observed in bloom in May. After the emergence in May of the first brood of the Karner Blue that was also observed, the team returned to the site where spotted knapweed, Centaurea maculosa Lam., was then in full bloom, serving as a nectar source. Approximately 2,000-3,000 Karner Blue butterflies were observed in a 1/3 acre (0.1349 ha) area. The calculated DD50 was 1331.

It was the Poweshiek Skipperling, O. poweshiek (Fig. 1) that led the author to the realization that degree-days would be a valuable tool in determining the emergence date of a species especially in historically-active sites. On 10 July 1997, the author traveled to an historically-active site discovered by Richard W. Holzman (Past-President, Michigan Nature Association (MNA), personal

Figure 2. Resident butterfly: Karner Blue, Plebejus melissa samuelis (Nabokov, 1944), 7 July 2007, Gates Road power-line, Reynolds Township, Montcalm County, MI, Man-istee National Forest (photo courtesy of Todd Smith).


communication) at Rattalee Lake Road, Oakland County, where the author’s first collection of the species occurred. In subsequent years as a member of MNA voucher specimens were collected on the MNA Clifford R. & Calla C. Burr Memorial Plant Preserve, contiguous to the Rattalee Lake Road site. On 29 June 1998 the author collected a voucher specimen at the MNA Big Valley Plant Preserve, Buckhorn Lake fen in Oakland County for a new site record. Using the degree-day values accumulated, the author pursued the species at an earlier DD value. On 22 June 2000 with a DD50 value of 848 at the nearby Milford weather station, a voucher specimen was collected at the MNA Burr Plant Preserve for the earliest day of the year flight period and the lowest ac-curate DD50 value. This information allows for further explorations at other potential sites and reevaluation of historically-active sites for the continued presence of this Michigan Natural Features Inventory (MNFI) Special Concern species. Degree-days provide information on the optimum early day to start a survey for a species and gives future explorations a more likely chance of specimen capture.

Degree Day counts could also be used to standardize counts of butterflies on one day per year, such as the Fourth of July Butterfly Counts. These kinds of surveys are extensive, however, the data collected from such surveys have not been widely utilized because of the difficulty in interpreting counts in which butterfly abundances appear to fluctuate. Fluctuations are due in part to the timing of when counts were made relative to phenology that is determined in part by growing degree days. The degree day approach could be used to standardize these long-term records across years and make the results useful.

Examples of the Michigan flight dynamics tend to fall into four categories: Resident, Migrant, Resident/Migrant, and Stray. The Michigan species are la-beled as such in Table 1. Listed by category are examples of these flight dynam-ics. Resident species include: Poweshiek Skipperling (Fig. 1); Karner Blue (Fig. 2); Dukes’ Skipper, Euphyes dukesi dukesi (Lindsey, 1923) (Hesperidae) (Fig. 3); and Swamp Metalmark, Calephelis muticum McAlpine, 1937 (Riodinidae) (Fig. 4). Migrant species include: Snout Butterfly, Libytheana carinenta bach-manii (Kirtland, 1851) (Nymphalidae) (Fig. 5); Fiery Skipper, Hylephila phyleus phyleus (Drury, 1773) (Hesperidae) (Fig. 6); and Painted Lady, Vanessa cardui (Linnaeus, 1758) (Nymphalidae). Migrant/Resident species include: Monarch, Danaus plexippus plexippus (Linnaeus, 1758) (Nymphalidae) (Fig. 7); Cloudless Sulphur, Phoebis sennae sennae = eubule (Linnaeus, 1767) (Pieridae) (Fig. 8); and Little Sulphur, Pyrisitia lisa lisa (Boisduval & Le Conte, 1830) (Pieridae). Strays include: Ocola Skipper, Panoquina ocola ocola (W. H. Edwards, 1863) (Hesperidae) (Fig. 9); Old World Swallowtail, Papilio machaon hudsonianus A. Clark; 1932 (Papilionidae); White-M Hairstreak, Parrhasius m-album (Boisduval & Le Conte, 1833) (Lycaenidae) (Fig. 10); and Goatweed Butterfly; Anaea andria Scudder, 1875 (Nymphalidae).

CONCLUSIONSI invite interested lepidopterists to use the information provided to de-

termine prior early flight period emergence/dispersal for Michigan butterflies and skippers, and eventually for the moth population. Of course the exact site location, nearby meteorological station, recorded degree-day value, and other pertinent data should be recorded for the species for that date and the new early date and DD50 should be published and/or provided to the Michigan Lepidoptera Survey for inclusion in the primary (Michigan listed species) and secondary (all Michigan species) databases.


Figure 3. Resident skipper: Dukes’ Skipper, Euphyes dukesi dukesi (Lindsey, 1923), 15 July 2005, Liberty Road fen, Liberty Township, Jackson County, MI (photo courtesy of Chris Rickards).

Figure 4. Resident butterfly: Swamp Metalmark, Calephelis muticum McAlpine, 1937, 13 July 2005, Lost Nation State Game Area, Jefferson Township, Hillsdale County, MI (photo courtesy of Dwayne R. Badgero).


Figure 5. Migrant butterfly: Snout Butterfly, Libytheana carinenta bachmanii (Kirt-land, 1851), 23 July 2005, Crosswinds Marsh, Sumpter Twonship, Wayne County, MI (photo courtesy of Chris Rickards).

Figure 6. Migrant skipper: Fiery Skipper, Hylephila phyleus phyleus (Drury, 1773), 20 August 2005, Gateway Garden, University of Michigan’s Matthaei Botanical Gardens, Superior Township, Washtenaw County, MI (photo courtesy of Chris Rickards).


Figure 7. Migrant/Resident butterfly: Monarch, Danaus plexippus plexippus (Linneaus, 1758), 15 July 2007, Greene Road, Moran Township, Mackinac County, MI (photo courtesy of Todd Smith).

Figure 8. Migrant/Resident butterfly: Cloudless Sulphur, Phoebis sennae sennae = eu-bule (Linneaus, 1767), 12 August 2006, Crosswinds Marsh, Sumpter Township, Wayne County, MI (photo courtesy of Chris Rickards).


Figure 9. Stray skipper: Ocola Skipper, Panoquina ocola ocola (W. H. Edwards, 1863), collected by Roger Kuhlman, 2 October 2005, Petersburg State Game Area, Summer-field Township, Monroe County, MI (photo courtesy of Ronald Priest and adapted by Owen A. Perkins).

Figure 10. Stray butterfly: White-M Hairstreak, Parrhasius m-album (Boisduval & Le Conte, 1833), collected by John Taggart, 12 August 1964, Duck Lake State Park, Fruit-land Township, Muskegon County, MI (photo by Owen A. Perkins).


Further investigation as to the peak flight periods and the life history degree-day values for the various species is encouraged.

ACKNOWLEDGMENTSI thank the following for their work in this field leading me to assemble

for use by others the valuable data related to degree-days and lepidoptera emergence: Harry D. King, who led the Michigan contingent into using the DD50 methodology; Robert D. Kriegel, who has the analytical mind and back-ground in this particular use of scientific data and who has gone before me in producing a paper showing the model for use of DD50 values for lepidoptera; and my colleague Mogens C. “Mo” Nielsen, with whom I have communicated for well over 50 years, sharing collecting experiences, information, and many hours in the field.

I thank my recent special associates and companions in the field, Dwayne R. Badgero, who has spent hours with me pursuing the documentation of Michi-gan species, and the historical and new habitats of Dukes’ Skipper; and Roger Kuhlman, whose Greater Washtenaw County Butterfly Survey Homepage website provides large amounts of data pertaining to the early and late dates.

I appreciate encouragement of Gary L. Parsons, Collections Manager, A. J. Cook Arthropod Research Collection at M.S.U., for me to contribute to the collection by recording the data of the Michigan lepidoptera for the MLS data-base. I also thank the professional reviews of the manuscript and supportive comments by Nick Haddad, North Carolina State University, and especially Therese M. Poland, USDA Forest Service, for her valuable editorial comments and agreement pertaining to publication in color.

I would like also to let it be known that all those who have collected and provided data to the MLS as well as those listed in Table 1 for whom early date specimens give validity to the project are most appreciated.

LITERATURE CITEDAndresen, J. A., and J. R. Harman. 1994. Springtime freezes in western Lower Michi-

gan: Climatology and trends, Michigan State University Extension. MSU Extension Vegetable Bulletins - AF089402.

Baskerville, G. L., and P. Emin. 1969. Rapid estimation of heat accumulation from maximum and minimum temperatures. Ecology 50: 514-517.

(MSU AWO) Michigan State University Agricultural Weather Office. 2008. Growing degree day accumulations. Available at: <http://www.agweather.geo.msu.edu/AWO/Current/report.asp?fileid=degreeday>

Nielsen, M. C. 1999. Michigan Butterflies and Skippers. Michigan State University Extension. East Lansing, Michigan.

(NOAA) National Oceanic and Atmospheric Administration. 2005. Quality controlled local climatological data. National Climatic Data Center, Asheville, North Carolina. Available at: <http://cdo.ncdc.noaa.gov/qclcd/QCLCD>

Pelham, J. P. 2008. A Catalogue of the Butterflies of the United States and Canada with a complete bibliography of the descriptive and systematic literature. J. Res. Lep. vol 40.


1Department of Biology, Drake University, Des Moines, Iowa 50311. 2 Department of Environmental Science and Policy, Drake University, Des Moines, Iowa 50311. * Corresponding author: (e-mail: [email protected]).

GREAT DIVERSITY OF INSECT FLORAL ASSOCIATES MAY PARTIALLY EXPLAIN ECOLOGICAL SUCCESS OF POISON IVY

(TOXICODENDRON RADICANS SUBSP. NEGUNDO [GREENE] GILLIS, ANACARDIACEAE)

David S. Senchina1* and Keith S. Summerville2

ABSTRACTLittle is known about insect floral associates of poison ivy (Toxicodendron

radicans, Anacardiaceae), despite the species’ ubiquity and importance in nature and society. Poison ivy’s pollination syndrome and results from prior studies suggest that the plant is not specialized for any particular pollinator type; how-ever, a systematic survey exploring this hypothesis has been lacking. For this study, insect floral associates of Toxicodendron radicans subsp. negundo from a central Iowa site were observed during the flowering season of 2005. Thirty-seven distinct insect floral associates were observed: 8 coleopterans (beetles), 7 dipterans (flies), 2 hemipterans (true bugs), 19 hymenopterans (ants, bees, wasps), and 1 lepidopteran (butterfly). Hymenopterans appeared to be the most important contributors to poison ivy pollination on a per species basis; however, coleopterans and dipterans were also frequent associates. Poison ivy’s ability to utilize a diverse assemblage of insect pollinators may partially explain its ecological success in varied habitats.

____________________

Toxicodendron radicans [L.] Kuntze (eastern poison ivy) is found com-monly throughout the eastern half of the United States. Several subspecies are recognized (Gillis 1971). T. radicans is one of five members of the poison-ous Toxicodendron P. Mill sect. Toxicodendron in North America, which also includes T. rydbergii [Small ex. Rydb.] Greene (Rydberg’s or western poison ivy), T. pubescens P. Mill. (syn. T. toxicarium Gillis, Atlantic poison oak), T. diversilobum [Torr. & Gray] Greene (Pacific poison oak), and T. vernix [L.] Kuntze (poison sumac). On the American supercontinent, members of this clade span the continental United States and are found in Canada and Mexico; other Toxicodendron spp. stretch into South America (McNair 1925, Barkley 1937, Gillis 1971).

Insect-host interactions have been described for poison ivy previously (reviewed in Senchina 2008a). These have focused on herbivory (Howden et al. 1951; Steyskal 1951; Hicks 1952, 1955), insect-host interactions (Gillis 1971, Tietz 1972, Robinson et al. 2006), or simply ecological association (Isaak and Honda 2002). Due to known natural histories, some of these associations may be predicted to be floral associations (specifically some involving the Hymenoptera and Lepidoptera), though they are not always catalogued specifically as such. Many of the above observations do not relate to floral association and it is some-times unclear which species of Toxicodendron is being reported due to the use of common names alone or confusion regarding scientific names. Thus, on several fronts, better data is needed regarding insect interactions with T. radicans in general, and insect pollination of the species in particular.


An examination of the pollination syndrome of T. radicans suggests that it is not specialized towards one specific type of pollinator. The plant is dioecious (e.g., male and female flowers are borne on separate plants) though each contains reduced or rudimentary organs from the other (Gillis 1971). Poison ivy flowers are small, pentamerous cream-colored flowers arranged in thyrses that sprout from leaf axils and oftentimes hang from or are obscured by the leaves (when viewed from human height). The petals become more recurved as the flower ages. The corolla is flat and lacking depth, with the stigma slightly exert or parallel to the petal edge. The ovary of female flowers turns from white to black when it is no longer receptive to pollen (Gillis 1971). Inflorescences are variable in the number of flowers (Gillis 1971). For T. radicans, the inflorescences vary from a couple to several cm in length.

Pacific poison oak (T. diversilobum) has been used as the model for floral anatomy in this genus. The pollen is bright yellow and produced only by male flowers despite the presence of rudimentary stamens in female flowers (Copeland and Doyel 1940). Nectar is secreted from the disk epidermis (Copeland and Doyel 1940, Gillis 1971); however, to the best of our knowledge, no one has reported whether or not nectar is produced by male flowers of T. radicans specifically. Investigations of closely related genera may provide some insight. Rhus is a genus of nonpoisonous sumacs closely related to Toxicodendron (Yi et al. 2004). Wheeler (2001) reports that both male and female flowers of staghorn sumac (Rhus hirta [L.] Sudworth) produce nectar, though the males produce much less. Young (1972) states that both male-sterile and hermaphroditic flowers in the gynodioecious Rhus integrifolia (Nutt.) Benth. & Hook f. ex. Brewer & S. Wats and Rhus ovata S. Wats produced nectar. For the more distantly-related andromonoecious cashew plant (Anacardium occidentale L.), both the male and hermaphrodite flowers produced nectar though the hermaphrodite flowers produced significantly more (Wunnachit et al. 1992). These results from other genera suggest that Toxicodendron spp. likely produce nectar in both male and female flowers, but that production in male flowers is likely reduced compared to female flowers. Historically, botanists have commented that Toxicodendron flowers emit a scent similar to lily-of-the-valley (Miller and Martyn 1809) or jasmine and hyacinth (McNair 1921). Others refute this (Gillis 1971). Given the well-characterized morphological plasticity of the genus, scent too may be a variable trait.

Little is known about the pollination ecology of poison ivy despite its ubiquity and importance in nature and human society. In his monograph on Toxicodendron spp., Gillis (1971) gives only Apis mellifera L. (Hymenoptera: Apidae, the honeybee) specifically as a pollinator of poison ivy, and adds, “Other visitors that appear to transport pollen include other Hymenoptera and several Coleoptera.” Poison ivy pollen is nonallergenic and is a major component of honey in Iowa (Pammel and King 1930) as well as other states (Lieux 1981). More recently, Kraemer and Favi (2005) have identified the blue orchard or orchard mason bee, Osmia lignaria Say (Hymenoptera: Apoidea: Megachili-dae), as another pollinator. Mulligan and Junkins (1977) named several other Hymenopteran floral associates. Senchina (2005, 2008b) described specific behaviors from Coleoptera and Lepidoptera. Table 1 gives an overview of all purported poison ivy pollinators and nectar-seekers known prior to the pres-ent investigation, but does not include taxa that are merely listed (or implied) as floral associates/visitors. No systematic survey of insect floral associates of poison ivy has yet been conducted, and it is unknown whether other insect orders (such as Diptera and Hemiptera) are also important floral associates.

The objective of this study was to better characterize insect floral associ-ates of poison ivy by cataloguing floral visitors to a Midwestern population of Toxicodendron radicans subsp. negundo (Greene) Gillis. Findings from this investigation were combined with previous data to obtain a better understand-ing of the ecology of poison ivy.


MATERIALS AND METHODSStudy Populations. All observations were conducted at East River Valley

Access (ERVA) in Ames, IA. ERVA is a mixed ash (Fraxinus spp. L.: Oleaceae)/elm (Ulmus spp. L.: Ulmaceae) floodplain forest with some linden (Tilia ameri-cana L.: Tiliaceae), hackberry (Celtis occidentalis L.: Ulmaceae), maple (Acer spp. L: Aceraceae/Sapindaceae), and oak (Quercus spp. L.: Fagaceae). Prevalent shrub- and herb-layer species (intermingled with T. radicans) include Virginia creeper (Parthenocissus quinquefolia [L.] Planchon: Vitaceae), frost or river grape (Vitis riparia Michx.: Vitaceae), gooseberry (Ribes missouriense Nutt.: Grossulariaceae), raspberry (Rubus sp. L.: Rosaceae), wild parsnip (Pastinaca sativa L.: Apiaceae), and various grasses. The site experienced a severe and prolonged flood in 1993 from which the flora has been recovering.

Toxicodendron radicans subsp. negundo is the only poison ivy species inhabiting this site and this region of Iowa (Eilers and Roosa 1994). ERVA includes both climbing and nonclimbing individuals, as well as populations on the forest edge and the forest interior. For this particular site, climbing individuals were found only in the forest interior and nonclimbing individu-als only along the forest edge. Nonclimbing individuals constituted the bulk of the sample and were clustered in three major groups. Group 1 was east-facing and approximately 18 × 4 m in size. Group 2 was directly 16 m west of Group 1, west-facing (i.e., on the other side of a narrow strip of forest), and approximately 24 × 4 m in size. Group 3 was directly 48 m north of Group 2, on the same west-facing forest edge, and approximately 38 × 3 m in size. Group 1 is bordered by a footpath to the east side and vegetation on all other sides, whereas Groups 2 and 3 are bordered by a footpath to the west side and vegetation on all other sides. Younger and shorter individuals tend to occupy areas closer to the footpath, whereas older and taller individuals tend to occupy areas closer to the forest interior. Consequently, the number of flowering in-dividuals increased with distance away from the footpath (Toxicodendron spp. do not flower until their third year [Gillis, 1971]). From observations made in previous field seasons, it can be noted that approximately equal numbers of males and females inhabit the site. Some general measurements were taken during mid-July 2008 to determine population characteristics. Within the area specified for Group 1, poison ivy plants were clustered in dense patches, occupying approximately 50% of the total area. Their average spacing within these patches was about 25 cm, with an average height of 60 cm. In contrast, within the areas specified for Groups 2 and 3, poison ivy plants were more or less contiguous and occupied approximately 80% of the total area and were variably spaced 10 cm (closer to the footpath) to 20 cm (closer to the forest interior), with an average height of 65 cm. These differences may be the result of sunlight exposure (east- vs. west-facing). Hundreds of individuals were thus included in these areas. Only two nonclimbing individuals could be located for inclusion in the study (northwest of the three nonclimbing groups in a contiguous portion of the forest). Both climbing and nonclimbing individuals and both genders were included in the study. We did not distinguish between visits to male vs. female plants or climbing vs. nonclimbing in our records.

Observations and Specimen Identification. All field observations were conducted daily from 5 to 21 June 2005 by the same lone observer except for on days of continuous rain (8, 10, and 20 June). These dates represent the first and last days in which poison ivy was found flowering. In total, over 50 hrs of obser-vations were amassed. Each observational period lasted a minimum of 1 hour (20 total observational sessions, average = 1.8 hrs, range = 1-5 hrs). The time of day during which observations were conducted was varied to include morning (no earlier than 3 hrs. after sunrise), afternoon, and evening hours (no later than 30 min before sunset), though the bulk of sessions (13 out of 20) were conducted in the afternoon as this appeared to be the time of greatest pollinator activity.


The goal during observation was to identify as many floral visitors as pos-sible; since there was a lone observer and oftentimes multiple, simultaneous pollinators, it was difficult to record factors such as number of flowers visited or length of visit for each individual insect (though this was attempted). An Olympus D-540 camera was used to photograph or video insect floral associates. For this investigation, we opted not to collect insects due to the allergenic nature of the plant and the stability (Bogue 1894) of its allergenic element (urushiol). Photographs were compared to specimens collected throughout Iowa from the Drake University and Iowa State University entomological collections and also print and Internet resources (Yanega 1996, Borror and White 2004, Tripplehorn and Johnson 2006).

RESULTSInsects from several different orders were found to be floral associates

at this study site (Table 2). In total, we observed 37 distinct insects that were identifiable to the level of family: 8 coleopeterans, 7 dipterans, 2 hemipter-ans, 19 hymenopterans, and 1 lepidopteran. Coleopteran floral associates included 6 members of Cerambycidae (the long-horned beetles), 1 member of Cantharidae (soldier beetles), and 1 member of Cleridae (checkered beetles). Dipteran floral associates included 4 members of Syrphidae (hoverflies) and 1 each from Anthomyiidae, Asilidae (robber flies), and Bombyliidae (bee flies). Two hemipteran floral associates from the Lygaeidae (seed, milkweed, or cinch bugs) were observed. Hymenopteran floral associates were the most abundant and diverse on both individual insect visitor and total insect species levels. We recorded flower visits from 6 members of Andrenidae (mining bees), 4 members of Apidae (bumble bees and honeybees), 1 member of Chalcidoidea (parasitoid wasps), 3 members of Halictidae (sweat bees), 2 members of Sphecidae (mud-daubers and sandwasps), and 3 members of Vespidae (paper and potter wasps). Only one lepidopteran, the summer azure, Celastrina neglecta W. H. Edwards (Lycaenidae, the gossamer-winged butterflies), was observed; a more thorough treatment of this insect’s association with poison ivy may be found elsewhere (Senchina 2008b).

On a per species basis, the greatest diversity of insect floral associates was observed at the beginning of the observational period (specifically 8 and 9 June). Similarly, the total number of insects seen visiting poison ivy flowers on a per day basis was greatest at the beginning of June and subsequently dwindled. By mid-June, the same species observed associating with poison ivy had shifted toward servicing neighboring plants, mainly wild parsnip and white clover (Trifolium repens L.; Fabaceae).

DISCUSSIONOnly a few hymenopteran and coleopteran species were known to be T.

radicans pollinators or nectar-seekers prior to this study (Table 1). Here we report findings from the first systematic survey of insect floral associates of poison ivy (Table 2), conducted during the 2005 field season. To the best of our knowledge, this study is the first to report the existence of both dipteran and hemipteran floral associates for this species, and supplies further infor-mation on additional coleopteran and lepidopteran floral associates as well as hymenopteran floral associates other than the honeybee or blue orchard bee. Together, Tables 1 and 2 represent the currently known catalogue of insect floral associates of Toxicodendron radicans. Data on beetle floral associates at the same site is also available from 2004 (Senchina 2005). Four of the five beetle floral associates witnessed in 2004 were also observed in 2005.

From the present data and known life histories, it appears that Hymenoptera may be the most important order of insects for poison ivy pollination. Thus data


ble

1: K

now

n po

ison

ivy

polli

nato

rs a

nd n

ecta

r-se

eker

s pr

ior t

o th

is s

tudy

. O

rder

Fa

mily

Sp

ecie

s St

udy

Cole

opte

ra

Cant

hari

dae

Cha

ulio

gnat

hus

mar

gina

tus (

Fabr

iciu

s)

Senc

hina

, 200

5Co

leop

tera

Ca

ntha

rida

e C

haul

iogn

athu

s pe

nnsy

lvan

icus

(DeG

eer)

Se

nchi

na, 2

005

Cole

opte

ra

Cera

mby

cida

e An

alep

tura

line

ola

(Say

) Se

nchi

na, 2

005

Cole

opte

ra

Cera

mby

cida

e St

rang

alia

acu

min

ata

(Oliv

ier)

Se

nchi

na, 2

005

Cole

opte

ra

Cler

idae

E

nocl

erus

ros

mar

us (S

ay)*

Se

nchi

na, 2

005

Hym

enop

tera

An

dren

dida

e An

dren

a sp

p. F

abri

cius

M

ullig

an &

Jun

kins

, 197

7H

ymen

opte

ra

Apid

ae

Apis

mel

lifer

a L.

G

illis

, 197

1H

ymen

opte

ra

Hal

iciti

dae

Agap

oste

mon

vir

isce

ns (F

abri

cius

) M

ullig

an &

Jun

kins

, 197

7H

ymen

opte

ra

Hal

iciti

dae

Augo

chlo

ra p

ura

(Say

) M

ullig

an &

Jun

kins

, 197

7H

ymen

opte

ra

Hal

iciti

dae

Lasi

oglo

ssum

spp

. Cur

tis

Mul

ligan

& J

unki

ns, 1

977

Hym

enop

tera

M

egac

hilid

ae

Osm

ia li

gnar

ia S

ay

Kra

emer

& F

avi,

2005

Lepi

dopt

era

Lyca

enid

ae

Cel

astr

ina

negl

ecta

(W. H

. Edw

ards

) Se

nchi

na, 2

008b

* =

Thi

s in

sect

was

pre

viou

sly

iden

tified

as

Than

asim

us d

ubiu

s but

is m

ore

likel

y E

nocl

erus

ros

mar

us (D

r. Jo

nath

an M

awds

ley,

U.S

. Fis

h &

W

ildlif

e Se

rvic

e, p

erso

nal c

omm

unic

atio

n, O

ctob

er 2

005)

.


Table 2: Insect floral associates of poison ivy (Toxicodendron radicans subsp. negundo) observed during the 2005 flowering season at a site in Ames, Iowa. Order Family Species # of Days Observed

Coleoptera Cantharidae Chauliognathus pennsylvanicus (DeGeer) 6Coleoptera Cerambycidae Analeptura lineola (Say) 1Coleoptera Cerambycidae Euderces picipes (Fabricius) 1Coleoptera Cerambycidae Molorchus sp. Fabricius 1Coleoptera Cerambycidae Strangalia acuminata (Olivier) 5Coleoptera Cerambycidae Unidentified cerambycid 1 1Coleoptera Cerambycidae Unidentified cerambycid 2 1Coleoptera Cleridae Enoclerus rosmarus (Say) 1Diptera Anthomyiidae Unidentified anthomyiid 1Diptera Asilidae Laphria sp. 6Diptera Bombylliidae Anthrax analis Say 1Diptera Syrphidae Eupeodes americanus (Weidemann)* 1Diptera Syrphidae Unidentified syrphid 1 8Diptera Syrphidae Unidentified syrphid 2 1Diptera Syrphidae Unidentified syrphid 3 1Hemiptera Lygaeidae Lygaeus kalmii Stål 1Hemiptera Lygaeidae Unidentified lygaeid 1 1Hymenoptera Andrenidae Andrena sp. Fabricius 2Hymenoptera Andrenidae Unidentified andrenid 1 3Hymenoptera Andrenidae Unidentified andrenid 2 5Hymenoptera Andrenidae Unidentified andrenid 3 5Hymenoptera Andrenidae Unidentified andrenid 4 1Hymenoptera Andrenidae Unidentified andrenid 5 1Hymenoptera Apidae Apis mellifera L. 3Hymenoptera Apidae Bombus fervidus (Fabricius) 2Hymenoptera Apidae Xylocopa sp. 1 Latreille 1Hymenoptera Apidae Xylocopa sp. 2 Latreille 1Hymenoptera Chalcidoidea Unidentified chalcidoid 1 3Hymenoptera Halictidae Agapostemon virescens (Fabricius) 3Hymenoptera Halictidae Unidentified halictid 1 1Hymenoptera Halictidae Unidentified halictid 2 1Hymenoptera Sphecidae Sceliphron caementarium (Drury) 1Hymenoptera Sphecidae Unidentified sphecid 1 1Hymenoptera Vespidae Eumenes fraternus Say 2Hymenoptera Vespidae Vespula sp. 1 Thomson 6Hymenoptera Vespidae Vespula sp. 2 Thomson 1Lepidoptera Lycaenidae Celastrina neglecta (W. H. Edwards) 3

* = This species has alternatively been classified as Metasyrphus americanus Weide-mann.


from this study may be useful to beekeepers or other individuals in the honey production business, as the honeybee as well as several other bee species were found to be common pollinators of poison ivy. Coleopterans, dipterans, hemipterans, and lepidopterans might also play important roles. The degree to which these different insects efficiently move poison ivy pollen from one flower to another likely varies tremendously. For example, Chauliognathus spp. (Coleoptera: Cantharidae) as well as longhorned beetles (Coleoptera: Cerambycidae) are known to be pollenivores of Toxicodendron spp., whilst anthomyiid, asilid, and syrphid flies (Diptera) are also pollen-feeding (McAlpine 1987, MacRae and Rice 2007). During the process of feeding, they may only tangentially transfer pollen from one plant to another. In contrast, the activi-ties of the hymenopterans (such as A. mellifera and O. lignaria), who receive nectar rewards from poison ivy, may be considered much more significant to pollination from the perspective of the plant. Further research may help define a relative continuum by which pollen transfer efficiency among these floral associates could be compared.

Poison ivy populations are widely found throughout multiple habitat types, such as fields, forests, urban areas, and waste places (Gillis 1971). Its ecologi-cal success may be partially explained by the pollinator plasticity documented here: because it is not reliant on a single species or single order of insects for pollination, poison ivy is capable of reproducing in multiple ecosystem types, and may be capable of recruiting novel pollinator species when it spreads into new habitats.

Importantly, we also discovered additional herbivores during our investiga-tion. One hemipteran (an unidentified species from Family Thyreocoridae, the negro bugs) was observed daily feeding off poison ivy leaves both in the 2004 and 2005 field seasons. A coleopteran (an unidentified species from Family Curculionidae, the weevils) was observed on a few days during the 2005 season. We occasionally observed one hymenopteran herbivore (an unidentified species from Family Tenthredinidae, the sawflies).

Beyond its empirical value to botanists, ecologists, entomologists, and natural history interpreters, data from this study may also have direct ap-plication for natural areas managers. In general, herbicides have failed to control unwanted poison ivy populations and alternative strategies have been sought. Natural areas managers are considering biological control agents such as fungi, arthropods, birds, and mammals (reviewed in Senchina [2008a] based on effectiveness, selectivity, practicality, and indirect or side effects). The identification of floral associates may offer some assistance. Floral as-sociates could be used as vectors for spreading a virus, bacterium, or fungal agent that is specific for poison ivy. Although in theory one could diminish the presence of a particular pollinator and thus adversely influence a plant’s reproductive capacity, this is not a very viable option for poison ivy because its pollinators are known to interact with several plant species. A more ten-able alternative may be to augment herbivore (such as those discussed in the preceding paragraph) or pollenivore populations. Certain members of the Cerambycidae, for example, may interact with Toxicodendron spp. in such a way that their low interindividual pollen transfer rate is outweighed by heavy pollen feeding and possible damage to adjacent reproductive structures, thus negatively impacting poison ivy reproduction overall. Controlled greenhouse and laboratory studies would be the first step in determining the feasibility of such endeavors. However, given how little research exists on this topic, it is likely that many more ecological associates for Toxicodendron spp. have yet to be identified, and more extensive field research is necessary to both catalogue and characterize such interactions.


LITERATURE CITEDBarkley, F. A. 1937. A monographic study of Rhus and its immediate allies in North and

Central America, including the West Indies. Ann. Miss. Bot. Gar. 24: 265-299.Bogue, E. E. 1894. The durability of the poisonous property of poison ivy, Rhus radicans

L. (Rhus toxicodendron L.). Science 23(581): 163.Borror, D. J., and R. E. White. 2004. Peterson field guide to insects. Houghton Mifflin,

New York.Copeland, H. F., and B. E. Doyel. 1940. Some features of the structure of Toxicodendron

diversiloba. Am. J. Bot. 27: 932-939.Eilers, L. J., and D. M. Roosa. 1994. The vascular plants of Iowa: an annotated checklist

and natural history. University of Iowa Press, Iowa City.Gillis, W. T. 1971. The systematics and ecology of poison-ivy and the poison oaks (Toxico-

dendron, Anacardiaceae). Rhodora 73(793-796):72-159,161-237,370-443,465-540.Hicks, S. D. 1952. Another insect feeding on Rhus of the Toxicodendron section. Coleopt.

Bull. 6(2): 29.Hicks, S. D. 1955. A dermestid in poison ivy. Coleopt. Bull. 9(1): 9.Howden, H. F., A. T. Howden, and P. O. Ritcher. 1951. Insects feeding on poison oak (Rhus

toxicodendron L.). Coleopt. Bull. 5(2): 17-19.Isaak, M., and J. Y. Honda. 2002. Preliminary insect survey on western poison oak

(Toxicodendron diversilobum (Torrey & Gray)), coyote bush (Baccharis pilularis De Condolle), and toyon (Heteromeles arbutifolia (Lindley)) in the Santa Cruz mountains, California. Pan-Pac Entomol. 78: 289-306.

Kraemer, M. E., and F. D. Favi. 2005. Flower phenology and pollen choice of Osmia lignaria (Hymenoptera: Megachilidae) in Central Virginia. Environ. Entomol. 34: 1593-1605.

Lieux, M. H. 1981. An analysis of Mississippi (U.S.A.) honey: pollen, color, and moisture. Apidologie 12: 137-158.

MacRae , T. C., and M. E. Rice. 2007. Biological and distributional observations on North American Cerambycidae (Coleoptera). Coleopt. Bull. 61: 227-263.

McAlpine, J. F. 1987. Manual of nearctic Diptera, Monograph #28 (vol. 2). Agriculture Canada Research Branch, Ottawa.

McNair, J. B. 1921. The morphology and anatomy of Rhus diversiloba. Am. J. Bot. 8: 179-191.

McNair, J. B. 1925. The geographical distribution in North America of poison ivy (Rhus toxicodendron) and allies. Am. J. Bot. 12: 338-350.

Miller, P., and T. Martyn. 1809. The gardeners and botanists dictionary. (9th ed.). Law and Gilbert, London.

Mulligan, G. A., and B. E. Junkins. 1977. The biology of Canadian weeds. 23. Rhus radi-cans. Can. J. Plant Sci. 57: 515-523.

Pammel, L. H., and C. M. King. 1930. Honey plants of Iowa. Iowa State Geological Bul-letin #7. State of Iowa: Des Moines.

Robinson, G. S., P. R. Ackery, I. J. Kitching, G. W. Beccaloni, and L. M. Hernandez. 2006. HOSTS -- a database of the world’s lepidopteran hostplants. Available at: <http://www.nhm.ac.uk/research-curation/research/projects/hostplants/>

Senchina, D. S. 2005. Beetle interactions with poison ivy and poison oak (Toxicodendron P. Mill. sect. Toxicodendron, Anacardiaceae). Coleopt. Bull. 59: 328-334.

Senchina, D. S. 2008a. Fungal and animal associates of Toxicodendron spp. (Anacar-diaceae) in North America. Persp. Plant Ecol. Evol. Syst. 10: 197-216.


Senchina, D. S. 2008b. Summer Azure (Celastrina neglecta W. H. Edwards, Lycaenidae) nectaring on poison ivy (Toxicodendron radicans, Anacardiaceae). J. Lep. Soc. 62: 55-56.

Steyskal, G. 1951. Insects feeding on plants of the Toxicodendron section of the genus Rhus (poison ivy, oak, or sumac). Coleopt. Bull. 5(5/6): 75-77.

Tietz, H. M. 1972. An index to the described life histories, early stages and hosts of the macrolepidoptera of the continental United States and Canada (vol. II). A. C. Allyn for the Allyn Museum of Entomology, Sarasota, Florida.

Tripplehorn, C. A., and N. F. Johnson. 2006. Study of insects. Thompson, Brooks and Cole, New York.

Wheeler, A. G. 2001. Biology of the plant bugs (Hemiptera: Miridae): Pests, predators, opportunists. Cornell University Press: Ithaca.

Wunnachit, W., C. F. Jenner, and M. Sedgley. 1992. Floral and extrafloral nectar produc-tion in Anacardium occidentale L. (Anacardiaceae): An andromonoecious species. Int. J. Plant Sci. 153: 413-420.

Yanega, D. 1996. Field guide to Northeastern long-horned beetles (Coleoptera: Ceramby-cidae). Illinois Natural History Survey Manual #7. Illinois Department of Natural Resources, Champaign.

Yi, T., A. J. Miller, and J. Wen. 2004. Phylogenetic and biogeographic diversification of Rhus (Anacardiaceae) in the Northern Hemisphere. Mol. Phylogenet. Evol. 33: 861-879.

Young, D. A. 1972. The reproductive biology of Rhus integrifolia and Rhus ovata (Anac-ardiaceae). Evol 26: 406-414.


EFFICACY OF MORPHOLOGICAL CHARACTERS FOR DISTINGUISHING NYMPHS OF EPITHECA CYNOSURA AND

EPITHECA SPINIGERA (ODONATA: CORDULIIDAE) IN WISCONSIN

Robert B. DuBois1*, Kenneth J. Tennessen2, and Matthew S. Berg3

ABSTRACTAttempts to distinguish exuviae and last-instar nymphs of Epitheca

cynosura (Say) and Epitheca spinigera (Selys) (Odonata: Corduliidae) using lateral spine characters have proven to be unreliable, and recent use of setae counts on only one side of the prementum or one labial palp have led to confu-sion because these structures often hold unequal numbers of setae on the two sides of the same specimen. Based on exuviae of 67 reared E. cynosura and 55 reared E. spinigera from lakes throughout Wisconsin, we tested the efficacy of previously used character states for distinguishing these species and searched for new characters to improve the reliability of regional keys. The most reliable diagnostic character was the combined number of setae on both sides of the prementum and on both labial palps (≤ 35 – E. cynosura; ≥ 36 – E. spinigera), which correctly determined 96% of our specimens. For the small percentage of specimens that lie in the region of overlap in total setae number, we found that total exuviae length, cerci ÷ epiproct ratios of females, tubercle distance ÷ epiproct ratios of males, and the shape of the dorsal hook on segment 8 could be used to strengthen determinations.

____________________

Despite numerous revisions (Muttkowski 1911, 1915; Davis 1933; Kormon-dy 1959; Tennessen 1973), the difficult genus Epitheca (Odonata: Corduliidae) has caused much confusion in North America (May 1995). This genus is often referred to as Tetragoneuria by workers who relegate Epitheca princeps Hagen to the genus Epicordulia (see Walker 1966). Confusion about this genus has encompassed both the naming of species, with only half of the 20 names that have been referred to the genus still widely accepted today, and discriminating among species in both the adult (Donnelly 1992, 2001; Needham et al. 2000) and nymphal stages, of which the latter are our current focus. Four currently accepted species of Epitheca are known in Wisconsin, our focal region of study. E. princeps and E. canis (McLachlan) are readily distinguished as last-instar nymphs and exuviae. However, efforts to distinguish between the last-instar nymphs and exuviae of E. cynosura (Say) and E. spinigera (Selys) have had a long and vexing history.

Referring to E. cynosura and E. spinigera, Walker (1913) remarked that, “A careful comparison was made between the exuviae of these two species, but no differences could be detected between them, except that in spinigera the lateral abdominal appendages average slightly longer than those of cyno-sura. This difference, however, does not appear to be constant.” In contrast

1Department of Natural Resources, Bureau of Endangered Resources, Ecological Inven-tory and Monitoring Section, 1401 Tower Avenue, Superior, Wisconsin 54880. *Corresponding author: (e-mail: [email protected]).2125 North Oxford Street, Wautoma, Wisconsin 54982. 3Grantsburg High School, Biology Department, 480 East James Avenue, Grantsburg, Wisconsin 54840.


to Needham’s (1901) use of the divergence of the lateral spines of abdominal segment 9 to separate these species, Walker (1913) reported considerable variation in this character among individuals of the same species. Despite this finding, Needham and Heywood (1929) used divergence of abdominal segment 9 lateral spines as the single diagnostic character for separating E. cynosura from E. spinigera in their key to nymphs of Tetragoneuria. Garman (1927) found the nymphs of the species of Tetragoneuria in Connecticut to be so similar that he did not attempt to construct a key for their separation. In their key to nymphs of North American Anisoptera, Wright and Peterson (1944) noted that very little dependence can be placed on nymphal determi-nations in the genus Tetragoneuria. However, Needham and Westfall (1955) again used the divergence of the lateral spines on abdominal segment 9 in their key to separate the group that included E. cynosura from the group that included E. spinigera.

Kormondy (1959) conducted the first in-depth study of the taxonomy and systematics of Tetragoneuria that included the nymphs. He initiated the possible utility of counts of palpal and premental setae to distinguish nymphs of E. cynosura and E. spinigera, and noted that last instar nymphs of E. spinigera averaged larger than those of E. cynosura. However, he con-cluded that considerable variation in both counts of raptorial setae and in last-instar size made them unreliable characters for taxonomic purposes. In his summary remarks about these species, he stated that “no dependable morphological characters serve to separate individual last-instar larvae…” He noted that as populations, they differed significantly in a number of ways including the larger mean size of E. spinigera, the much higher growth rate of E. spinigera from the 11th to 12th instar, and the earlier seasonal appear-ance of E. spinigera.

Walker and Corbet (1975) used the number of premental setae (usually 10 or less for E. cynosura; usually 11 or more for E. spinigera) as the sole character to separate these species. In their diagnosis under E. spinigera, they reiterated that the relative length and direction of the lateral spines on segment 9 was not reliable, that overlap could be expected in both palpal and premental setae counts, and that E. spinigera averaged larger than E. cynosura, but only in part of the former’s range. In their on-line key, Bright and O’Brien (1998) separated the species using the number of premental setae (usually not less than 11 for E. spinigera; usually not more than 10 for E. cynosura), followed by the number of palpal setae (usually 7-8 for E. spinigera; usually 6 for E. cynosura). Needham et al. (2000) similarly used the number of premental setae (usually 11-12), followed by the number of palpal setae (usually 7) to separate E. spinigera from the group containing E. cynosura (premental setae usually 8-10; palpal setae usually 5-6). We tried to use these keys to distinguish last-instar nymphs and exuviae of E. cyno-sura and E. spinigera from waters in Wisconsin, but were uncertain about many determinations. Many specimens belonging to one of these species had 11 or more setae on one side of the prementum, but 10 or fewer setae on the other side, or at least 7 setae on one labial palp, but only 6 setae on the other. For some specimens, counting premental setae led to a different determination than counting palpal setae (e.g.,10 or fewer setae on both sides of the prementum indicating E. cynosura, but 7 or more setae on each palp indicating E. spinigera).

To make firmer recommendations for separating exuviae and last-instar nymphs of these species, we reared specimens of both species from numerous lakes in Wisconsin to form a database on which a number of morphological analy-ses could be performed. Our objectives were to determine if either premental or palpal setae counts, or a combination of the two, would reliably distinguish these species in Wisconsin, and to search for and evaluate other characters that could be used to separate them.


MATERIALS AND METHODSWe used only reared exuviae in our analyses to be certain of their identity.

Nymphs were collected after molting to F-0 and were reared to emergence in aquaria. Teneral adults were maintained alive in small cages for several days after emergence, then were soaked overnight in acetone, dried, and stored in standard Odonata envelopes. Exuviae were placed in individual vials of 70% ethanol. Each adult/exuvia association was given a unique accession number immediately after emergence to preclude any possibility of confusing the specimens.

Our dataset was comprised of exuviae of 55 E. spinigera and exuviae of 67 E. cynosura. All of the exuviae except four were reared by one of the authors, and the first author verified all determinations and made all counts and measure-ments. Not all exuviae were used in all analyses. Two exuviae of E. cynosura and one exuvia of E. spinigera lacked essential mouth parts and could not be used for raptorial setae counts. Three exuviae of E. cynosura and one exuvia of E. spinigera were not measured for total length because the heads were detached or missing. One exuvia of E. spinigera had a twisted epiproct and was not used in analyses involving that character. Exuviae of 6 E. spinigera and 1 E. cynosura were omitted from analyses involving abdomen dimensions, and exuviae of 3 male E. spinigera and 1 male E. cynosura were omitted from analyses involv-ing dimensions of the cerci and epiproct. All specimens are housed either in the Odonata Collection of the Wisconsin Department of Natural Resources (WDNR) in Superior, or in the private collection of KJT in Wautoma, Wisconsin.

Abdominal segments are designated by the letter “S” and the segment number (e.g., S9 = abdominal segment 9). Counts and measurements were done under magnification using either an ocular micrometer or millimeter rule, and all measurements are reported in mm. Counts of setae on both labial palps and on both sides of the prementum were done in dorsal view with the prementum pulled outward with a teasing needle. The most medially located premental setae were small, and care was taken to count all of them. Total lengths of exuviae were measured in dorsal view with a millimeter rule. We assessed the direction in which the tip of the dorsal hook on S8 pointed in lateral view rela-tive to the long axis of the body. The left lateral spine on S9 was measured for width at the base and length in strict dorsal view with an ocular micrometer. Also measured in dorsal view were the lengths of both of the lateral spines on S8, the lengths of the margins of S8 including the spines, and the lengths of both cerci and the epiproct. For the males, the length of the epiproct from the base to the distal margin of the ante-apical tubercles was also measured. The abdomen was measured in ventral view from the base of S1 to the apex of S10 to determine length, and across the width of S6 with the sclerite lightly depressed to determine maximum width.

Statistical analyses were performed using SigmaStat statistical software (SPSS 1997) with alpha set at 0.05. We used one way ANOVA (F), or Kruskal-Wallis one way ANOVA on ranks (H), to test for differences in setae numbers and total exuviae lengths among lakes and years. Pearson product moment correlation (r) was used to examine the strength of correlation between setae numbers and total exuviae lengths. The Chi-square test (χ2) was used to examine differences between species in the shape of the left lateral spine on S9 in dorsal view: categories were straight (1), and slightly incurved (2). Mann-Whitney Rank Sum tests (T) were used to examine differences between species in abdomen shape ratios. Statistical tests were not applied to other key characters because our goal was to assess the performance of character states in potential key couplets, not to determine statistical significance.

Material examined – WISCONSIN: BAYFIELD CO.: Sawdust Lake, 18 April and 5 May 2005, UTM 15 632537E 5159021N (all coordinates NAD83/WGS84), RBD (exuviae of 11 reared E. cynosura and exuviae of 3 reared E. spinigera);


Eau Claire River, 8 May 2007, UTM 15 615079E 5128789N, RBD (exuviae of 2 reared E. cynosura); BURNETT CO.: Dunham Lake, 23 April 2005, UTM 15 541560E 5067888N, MSB (exuviae of 17 reared E. cynosura); Clam River, 26 April 2007, UTM 15 553423E 5076631N, RBD (exuviae of 1 reared E. cynosura); pond, Memory Lake Campground, 22 May 2005, 10 April 2006, and 22 February 2007, UTM 15 524426E 5069876N, MSB (exuviae of 22 reared E. spinigera); Poquette Lake, 9 April 2005, UTM 15 571908E 5073126N, MSB (exuviae of 7 reared E. cynosura); DOUGLAS CO.: Amnicon Lake, 19 May 2006, UTM 15 571420E 5147735N, RBD (exuviae of 1 reared E. cynosura); Breitzman Lake, 7 June 2002 and 12 May 2003, UTM 15 560242E 5145202N, RBD (exuviae of 4 reared E. cynosura); Gander Lake, 4 June 2002, UTM 15 597475E 5145108N, RBD (exuviae of 2 reared E. cynosura); Rush Lake, 20 April 2007, UTM 15 610833E 5149561N, RBD (exuviae of 6 reared E. cynosura and exuviae of 6 reared E. spinigera); Cut-A-Way Dam, St. Croix River, 1 June 2005 and 16 May 2006, UTM 15 593329E 5128758N, RBD (exuviae of 2 reared E. cynosura and exuviae of 3 reared E. spinigera); Upper St. Croix Lake, 20 May 2005, UTM 15 593857E 5136942N, RBD (exuviae of 8 reared E. cynosura); EAU CLAIRE CO.: Half Moon Lake, 13 November 2007, UTM 15 616483E 4962190N, RBD (exuviae of 2 reared E. cynosura); LANGLADE CO.: Crooked Lake, 14 May 2003, UTM 16 327200E 5021066N, KJT (exuviae of 1 reared E. spinigera); ONEIDA CO.: Tomahawk Lake, Kemp Natural Resources Center, 14 June 2004, UTM 16 291966E 5079767N, RBD (exuviae of 1 reared E. spinigera); RICHLAND CO.: Cruson Slough, Wisconsin River, 21 April 2007, UTM 15 724367E 4785883N, W. A. Smith (exuviae of 4 reared E. cynosura); WAUSHARA CO.: Bohn Lake, 24 April 2006, UTM 16 304602E 4888113N, KJT (exuviae of 14 reared E. spini-gera); Meilke Lake, 12 May 2007, UTM 16 315894E 4877729N, KJT (exuviae of 5 reared E. spinigera).

RESULTSCombining the data on total number of palpal + premental setae (= total

setae) for the two species resulted in a bimodal curve (Fig. 1). Overlap occurred from 34 through 37 total setae, although few individuals were represented in this range. Approximately 95% of the E. cynosura had 35 or fewer total setae, whereas 96% of the E. spinigera had 36 or more total setae (Table 1). For E. cynosura, total setae counts did not differ among five lakes having the largest sample sizes (H = 2.616, df = 4, P = 0.624), and total setae number was not cor-related with total length of exuviae (r = -0.0673, N = 65, P = 0.594). However, for E. spinigera, total setae counts differed significantly among four lakes having the largest sample sizes (H = 9.427, df = 3, P = 0.024), and total setae number was positively correlated with total length of exuviae (r = 0.381, N = 54, P = 0.005).

Numbers of premental setae also resulted in a bimodal distribution, but substantial overlap occurred from 20 to 23 setae, with most overlap at 22 setae (Table 1). For E. cynosura, 86% of exuviae had 21 or fewer premental setae, with most exuviae having 19 to 21 setae. For E. spinigera, 96% of exuviae had 22 or more premental setae, with most exuviae having 22 to 25 setae. Exuviae of both species often had unequal numbers of setae on the two sides of their prementum; this occurred on 51% of E. cynosura and 56% of E. spinigera.

Substantial overlap in numbers of palpal setae between species occurred from 13 through 15 setae and the single highest percentage of both species had 14 setae (Table 1). Fifty nine percent of E. cynosura had 13 or fewer palpal setae, while only 4% of E. spinigera had that number. Twenty six percent of E. spinig-era had 15 or more palpal setae, while only 3% of E. cynosura had that number. Exuviae of both species sometimes had unequal numbers of setae on their two labial palps; this occurred on 34% of E. cynosura and 22% of E. spinigera.


Figu

re 1

. Num

bers

of t

otal

rapt

oria

l set

ae (l

eft +

righ

t pal

pals

plu

s le

ft +

righ

t pre

men

tals

) pos

sess

ed b

y re

ared

exu

viae

of

Epi

thec

a cy

nosu

ra a

nd E

. spi

nige

ra fr

om W

isco

nsin

.


Table 1. Frequency of exuviae with number of raptorial setae (left + right palpals, left + right prementals, and total) in reared Epitheca cynosura (n = 65) and E. spinigera (n = 54) from Wisconsin.

Palpals 12 13 14 15 15

E. cynosura 18 20 25 2 0E. spinigera 0 2 38 9 5

Prementals 17 18 19 20 21 22 23 24 25 26

E. cynosura 1 3 14 23 15 8 1 0 0 0E. spinigera 0 0 0 1 1 10 17 12 8 5

Total ≤31 32 33 34 35 36 37 38 39 ≥40

E. cynosura 6 11 18 17 10 2 1 0 0 0E. spinigera 0 0 0 1 1 9 16 10 7 10

Exuviae of E. spinigera averaged 22.9 mm in total length, while exuviae of E. cynosura averaged 21.9 mm in total length, but the overlap between species was substantial. Differences in mean exuviae length among lakes were evident for both species that could have been attributable to lake effects, inter-annual effects, or both. For E. cynosura total exuvia length was significantly different among four lakes sampled in 2005 (H = 13.195, df = 3, P = 0.004). Significant differences among lakes sampled in 2007 were found also for E. spinigera (F = 13.221, df = 3, P < 0.001). Our data allowed for only one direct test of an inter-annual effect on total exuviae length, at a pond at Memory Lake Campground in Burnett County, where E. spinigera were sampled in 2005, 2006, and 2007. Here we did not find evidence of an effect attributable to year (H = 1.566, df = 2, P = 0.457).

Female exuviae of E. spinigera had longer cerci than did female exuviae of E. cynosura. This was most clearly seen in the ratio obtained by dividing the mean length of both cerci by the length of the epiproct. Despite some overlap, the cerci ÷ epiproct ratio was < 0.75 for all female E. cynosura, and ≥ 0.75 for 90% of female E. spinigera (Table 2). We did not find a difference between species in the cerci ÷ epiproct ratio for male exuviae (T = 821.5, N = 65, P = 0.188).

The position of the ante-apical tubercles on the epiproct of male exuviae, expressed as the ratio of the distance from the base of the epiproct to the distal margin of the tubercle ÷ the total length of the epiproct, differed between species. The tubercles of E. cynosura were located more distally on the epiproct than those of E. spinigera (Table 2). However, the difference between species was slight and there was some overlap. Nearly all (98%) E. cynosura had a tubercle distance from base ÷ epiproct length ratio ≥ 0.66, whereas 73% of E. spinigera had values < 0.66.

The shape of the dorsal hook on S8, with respect to the direction in which the tip was pointed in strict lateral view, sometimes differed between species (Fig. 2). Often (55% of E. cynosura; 36% of E. spinigera), the tip of the dorsal hook on S8 pointed straight rearward in line with the long axis of the body. In these cases, the character would have had no diagnostic value. However, among the remainder, the tip of the hook pointed either slightly downward (ventrally – Fig. 2a) or slightly upward (dorsally – Fig. 2b) relative to a long-axis body line.


Table 2. Mean character ratios of reared exuviae of Epitheca cynosura and E. spinigera from Wisconsin (SE in parentheses).

♂ Base to ♀ Cerci/ tubercle/ Spine of 8/ Spine of 9 AbdomenSpecies epiproct epiproct margin of 8 W at base/L W at S6/L

E. cynosura 0.656 0.708 0.181 0.298 0.685 (0.00746) (0.00381) (0.00253) (0.00458) (0.00275)

E. spinigera 0.783 0.645 0.163 0.301 0.692 (0.00701) (0.00623) (0.00211) (0.00533) (0.00416)

Figure 2. Dorsum of abdomen in lateral view of exuviae of Epitheca spinigera (a) and E. cynosura (b); arrows indicate direction of S8 hook relative to long axis of body (repre-sented by horizontal line).


All cases in which the tip pointed slightly upward were E. cynosura, and 93% of cases in which the hook pointed slightly downward were E. spinigera.

The lateral spines on S8, relative to the length of the lateral edge of the segment including the spine, averaged longer on E. cynosura, but the mean difference in this ratio between species was slight (Table 2) and there was substantial overlap. The slenderness of the left lateral spine on S9, expressed as the ratio of width at the base ÷ length, did not differ between species (Table 2). The shape of the left lateral spine on S9 (straight vs. slightly incurved) also did not differ between species (χ2 = 1.002, df = 1, P = 0.317).

The shape of the abdomen (ratio of the maximum width at S6 ÷ total abdomen length) did not differ between exuviae of the two species (T = 3028.0, P = 0.229; Table 2). The maximum width of the abdomen at S6 was greater for E. spinigera (avg. = 8.6 mm) than for E. cynosura (avg. = 8.1 mm); however, this difference was attributable to the larger mean size of E. spinigera, not to a difference in shape, and there was substantial overlap in the character.

DISCUSSIONRaptorial setae counts reliably distinguished between reared exuviae

of E. cynosura and E. spinigera, but only if all premental setae, or preferably, total setae (all premental + all palpal setae) were counted. An optimal couplet using total setae counts (E. cynosura ≤ 35; E. spinigera ≥ 36) was most effective, and would have correctly determined 96% of the reared exuviae in our sample, with no substantial overlap at any single number of total setae. An optimal couplet using just premental setae counts (E. cynosura ≤ 21; E. spinigera ≥ 22) would have correctly determined 91% of the reared exuviae in our sample, with substantial overlap occurring only at 22 setae. The increase in efficacy gained by use of total setae counts over just premental setae counts is worthwhile and easily attained because palpal setae are easy to count, and the labial palps are exposed and readily visible when the setae on the prementum are counted. Pal-pal setae counts alone were unreliable for distinguishing between these species because there was considerable overlap at 14 setae, and the most reasonable diagnostic couplet using palpal setae (E. cynosura ≤ 13; E. spinigera ≥ 14) would erroneously determine 42% of E. cynosura as E. spinigera.

When setae counts are used, we urge that setae on both sides of the premen-tum and on both labial palps be counted because we frequently found unequal numbers of these setae on the two sides of exuviae of both species. Counting setae on both sides of both structures provides results that are less ambiguous and less confusing than employing counts on only one side, as has been the current practice (Walker and Corbet 1975, Bright and O’Brien 1999, Needham et al. 2000). For example, premental setae counts were used in these keys as either the sole character, or the primary character, to separate E. spinigera (11 or more setae on one side) from E. cynosura, or the group containing E. cynosura (10 or fewer setae on one side). Based on our data, this character would erroneously determine 14% of E. cynosura that have at least 11 setae on each side of the prementum, and would leave ambiguous another 25% of E. cynosura that have 11 setae on one side of the prementum. Thus, the character would correctly and unambiguously determine only 62% of E. cynosura. Adding palpal setae counts to the couplet as a secondary character (Bright and O’Brien 1999, Needham et al. 2000) did not reduce the confusion because a substantial proportion of both species have unequal numbers of palpal setae on their two sides.

We do not generally support use of total exuviae length as a diagnostic character to distinguish these species even though E. spinigera is often somewhat larger. Difficulties could occur because the area of overlap in length is extensive, and some lakes have small E. spinigera. Kormondy (1959) noted a north-south cline of last-instar size in Michigan, with smaller individuals in the southern


part of the state. Exuviae ≥ 24 mm in total length in Wisconsin are at least 12 times more likely to be E. spinigera than E. cynosura. However, only 38% of E. spinigera in our sample attained that size. If total setae counts of a last-instar nymph or exuviae are in the center of the region of overlap between species (35 or 36 setae), then a total length ≥ 24 mm would strongly indicate E. spinigera. The findings that total lengths of exuviae of both species varied among lakes, and that total lengths of E. spinigera were positively correlated with total setae counts, suggest that separating these species using setae counts will be most challenging with small E. spinigera that sometimes occur in certain lakes. Our inability to demonstrate an inter-annual effect on total exuviae length for E. spinigera over three years at just one site does not necessarily mean that this character does not vary among years in either species at other sites.

The finding that the cerci ÷ epiproct ratio differed significantly between female exuviae of these species was not unexpected because adult female E. spinigera have considerably longer cerci than do adult female E. cynosura. This ratio could have limited utility as an ancillary diagnostic character to separate female exuviae and last instar nymphs of these species (cerci at least 3/4 the length of the epiproct = E. spinigera; cerci less than 3/4 the length of the epiproct = E. cynosura). Although there is relatively little overlap between species, the cerci ÷ epiproct ratio has drawbacks that would reduce its diagnostic value in most circumstances including: 1) most specimens of both species are quite close to the 0.75 ratio cutoff value, requiring precise measurements to be made, 2) the two cerci on a single specimen may differ slightly in length, requiring that both be measured and averaged, which takes additional time, 3) the character can be applied to specimens of only one gender, and 4) reliability in excess of 95% can be achieved using the simpler character of total setae counts alone. However, in cases where last-instar nymphs and exuviae have total setae counts of 35 or 36, and total lengths that are < 24 mm, calculating the cerci ÷ epiproct ratios of females could aid in making determinations.

The finding that the position of the ante-apical tubercles on the epiprocts of male exuviae differs somewhat between species also has limited diagnostic value, because as with the previous character, the difference was slight (thus requiring exacting measurements), there was some overlap, the character can be applied to only part of the population, and a better diagnostic character exists. As with the cerci ÷ epiproct ratios of females, the tubercles ÷ epiproct ratios of males (< 0.66 = E. spinigera; ≥ 0.66 = E. cynosura) could be helpful to weigh into the identification process in cases where last-instar male nymphs and exuviae have total setae counts of 35 or 36, and total lengths < 24 mm.

We hesitate to recommend use of dorsal hook and lateral spine charac-ters to separate these species, even though some significant differences were found, because substantial variation occurred in these characters that could have been environmentally induced. The direction in which the tip of the dorsal hook on S8 points could cautiously be used as an ancillary character to reinforce determinations made using other characters. In the approximately 54% of the exuviae of the two species in our sample in which the tip of this hook did not point straight rearward in line with the long axis of the body, an upward point-ing tip strongly indicated E. cynosura and a downward pointing tip suggested E. spinigera, though somewhat less strongly. Although the lateral spine on S8 averaged significantly longer relative to the length of the segment margin on E. cynosura, the difference between the species was slight and there was considerable overlap, which reduced the diagnostic value of the character. Our assessments of the shape and dimensions of the left lateral spine on S9 led us to the same conclusion reached by Walker (1913) and Kormondy (1959), that there are no useful diagnostic attributes associated with this spine.

Contrary to our expectations, the shape of the abdomen was highly vari-able for each species, and it did not differ between species. Part of the reason


for this variation could have been associated with the difficulty, because of the various contorted shapes of exuviae, of depressing the sclerite on the venter of segment 6 in a consistent way to determine maximum width, and of obtaining a consistent value for total abdomen length.

RECOMMENDATIONSUse of total setae counts (E. cynosura ≤ 35; E. spinigera ≥ 36) is the most

effective way to separate exuviae and last-instar nymphs of these species. These counts correctly determined 96% of the reared exuviae in our sample, with no substantial overlap at any single number of total setae. For specimens in the center of the area of slight overlap (35 or 36 total setae), several ancillary characters can be used to reinforce determinations including: total length (if ≥ 24 mm – E. spinigera), cerci ÷ epiproct ratio for females (E. cynosura < 0.75; E. spinigera ≥ 0.75), distance to ante-apical tubercle ÷ epiproct for males (E. spinigera < 0.66; E. cynosura ≥ 0.66), and the direction the tip of the dorsal hook on segment 8 projects in lateral view (slightly upward – E. cynosura; slightly downward – E. spinigera). These recommendations are intended for application only with last-instar nymphs and exuviae and only within the region we sampled (Wisconsin). Extrapolation of these recommendations with younger nymphs or outside this region should be undertaken cautiously.

ACKNOWLEDGMENTSFunding for RBD was provided by the Bureau of Endangered Resources

of the Wisconsin Department of Natural Resources. We thank W. A. Smith for rearing 4 specimens of E. cynosura and J. M. Pleski for help in the field and with rearing.

LITERATURE CITEDBright, E., and M. F. O’Brien. 1998. Odonata larvae of Michigan: keys for, and notes on the

dragon- and damselfly larvae found in the state of Michigan. Key to mature larvae of Michigan Epitheca. Available at: <http://insects.ummz.lsa.umich.edu/michodo/test/Epitheca.htm>

Davis, W. T. 1933. Dragonflies of the genus Tetragoneuria. Bulletin of the Brooklyn En-tomological Society 28: 87-104.

Donnelly, T. W. 1992. Taxonomic problems (?) with Tetragoneuria. Argia 4: 11-12.Donnelly, N. 2001. Taxonomic problems with North American Odonata species – a last

appeal for information. Argia 13: 5-10.Garman, P. 1927. Guide to the insects of Connecticut. Part V, the Odonata or dragonflies

of Connecticut. Connecticut Geological and Natural History Bulletin 39. 331 pp.Kormondy, E. J. 1959. The systematics of Tetragoneuria, based on ecological, life history,

and morphological evidence (Odonata: Corduliidae). Miscellaneous Publications, Museum of Zoology, University of Michigan, No. 107. 79 pp.

May, M. L. 1995. The subgenus Tetragoneuria (Anisoptera: Corduliidae: Epitheca) in New Jersey. Bulletin of American Odonatology 2: 63-74.

Muttkowski, R. A. 1911. Studies in Tetragoneuria (Odonata). Bulletin of the Wisconsin Natural History Society 9: 91-134.

Muttkowski, R. A. 1915. Studies in Tetragoneuria (Odonata). II. Bulletin of the Wisconsin Natural History Society 13: 49-61.

Needham, J. G. 1901. Aquatic insects in the Adirondacks. Order Odonata. Bulletin of the New York State Museum 47: 381-612.


Needham, J. G. and H. B. Heywood. 1929. A handbook of the dragonflies of North America. Springfield, Illinois, C. C. Thomas. 378 pp.

Needham, J. G. and M. J. Westfall. 1955. A manual of dragonflies of North America (Anisoptera). Berkeley, University of California Press. 615 pp.

Needham, J. G., M. J. Westfall, and M. L. May. 2000. Dragonflies of North America. Gainesville, Florida, Scientific Publishers. 939 pp.

SPSS. 1997. SigmaStat for Windows, version 2.03. SPSS, Chicago.Tennessen, K. J. 1973. A preliminary report on the systematics of Tetragoneuria (Odonata:

Corduliidae) in the southeastern United States. M.S. Thesis, University of Florida, Gainesville.

Walker, E. M. 1913. New nymphs of Canadian Odonata. Can. Entomol. 45: 161-170.Walker, E. M. 1966. On the generic status of Tetragoneuria and Epicordulia (Odonata:

Corduliidae). Can. Entomol. 98: 897-902.Walker, E. M., and P. S. Corbet. 1975. The Odonata of Canada and Alaska. Vol. 3. Toronto,

Ont., University of Toronto Press. 307 pp.Wright, M., and A. Peterson. 1944. A key to the genera of anisopterous dragonfly nymphs

of the United States and,Canada (Odonata, suborder Anisoptera). Ohio J. Sci. 44: 151-161.


WHY ARE THERE SO FEW INSECT PREDATORS OF NUTS OF AMERICAN BEECH (FAGUS GRANDIFOLIA)?

Charles E. Williams1

ABSTRACTAmerican beech, Fagus grandifolia Ehrh., is a common nut-bearing tree

of eastern North America. Compared to other North American nut-bearing tree species of comparable geographic range, the nut-infesting insect fauna of American beech is species-poor: only the filbertworn, Cydia latiferreana (Wlsm.) (Lepidoptera: Tortricidae), infests nuts of American beech. Why are there so few insect predators of nuts of American beech? Using data from published studies, I explore two hypotheses that may help to explain the species-poor nut-infesting insect fauna of American beech. First, might chemical defense of beechnuts, and/or low nutritional value, restrict the number of insect predators that can exploit this food resource (unprofitable resource hypothesis)? Second, may spatial and temporal variability of beechnut mast crops limit colonization by nut-infesting insects because of the unpredictability of the resource (unpredictable resource hypothesis)? I found no strong evidence to suggest that chemical defense or low nutritional value was associated with the species-poor nut-infesting insect fauna of American beech. Yearly variability in nut crop size alone did not explain the low species richness of American beech compared to other tree species. Instead, I suggest that spatial and temporal unpredictability in production of sound versus incomplete beechnuts was an effective filter that limited colonization of beechnuts by insects. Moreover, the lone insect species able to successfully colonize beechnuts, C. latiferreana, is well adapted to resource unpredictability. Unlike specialist insect species that infest nuts of only 1 or 2 North American tree genera, C. latiferreana has a relatively broad host range and its mobile larvae can relocate to new resources when faced with food shortages.

____________________

Considerable attention has focused on the biogeographic relationships of herbivorous insects and their host plants, especially on factors that affect ac-cumulation of leaf-feeding insects by plants in ecological and evolutionary time (Strong, Jr. 1974, 1979, Blaustein et al. 1983, McCoy and Rey 1983, Strong, Jr. et al. 1984). Research has shown that accrual of leaf-feeding insects by plants is influenced by several factors including plant geographic range, architecture, and toxicity (Strong, Jr. et al. 1984 and references therein).

Little studied in a biogeographic context, fruits and seeds of plants may also support diverse communities of insects (Winston 1956, Andersen and New 1987). Evidence suggests that accrual of fruit- and seed-eating insects by plants is influenced by factors similar to those that affect accumulation of leaf-feeding insects. For example, Andersen and New (1987) found that host plant phylogeny and fruit morphology were important correlates of the distribution and abundance of seed-eating insects of fruits of Australian Eucalyptus, Lep-tospermum, and Casuarina.

American beech, Fagus grandifolia Ehrh., a common tree species of east-ern North America, is a member of the Fagaceae, a nut-bearing woody plant family whose fruits are extensively used as food by animals, especially insects

1Western Pennsylvania Conservancy, Allegheny Field Office, 40 W. Main Street, Ridg-way, PA 15853. (e-mail: [email protected]).


(Martin et al. 1951, Marquis et al. 1976, Williams 1989). However, compared to other nut-bearing tree species in North America, the nut-infesting insect fauna of American beech is decidedly species-poor (Williams 1989). Only one species, Cydia latiferreana (Wlsm.) (Lepidoptera: Tortricidae), the filbertworm, infests nuts of American beech. In contrast, the nut-infesting insect faunas of most other North American tree genera of comparable geographic range are substantially more species-rich (Table 1, Williams 1989), particularly the major fagaceous genera, Quercus (oaks) and Castanea (chestnuts, chinkapins) (plant nomenclature follows Gleason and Cronquist 1991). Similarly, nuts of European beech, Fagus silvatica L., are infested only by Cydia fagiglandana Zeller, a European relative of the filbertworm (Watt 1923, Nielsen 1977, Jensen 1985, Nilsson 1985).

Why are there so few insect predators of nuts of American beech? Using data from published studies, I explore two hypotheses that may help explain the species-poor nut-infesting insect fauna of American beech. First, might chemical defense of beechnuts, and/or low nutritional value, restrict the number of insect predators that can exploit this food resource (non-profitable resource hypothesis)? In particular, tannins, common chemical constituents of nuts of many fagaceous species, can influence feeding preferences in animals (Smallwood and Peters 1986) and can bind with proteins in the digestive tract, rendering them indigestible (Martin and Martin 1982). Second, may spatial and temporal variability of beechnut mast crops limit colonization by nut-infesting insects because of the unpredictability of the resource (unpredictable resource hypoth-esis)? Masting, the synchronous production of seed crops at irregular intervals (Silvertown 1980, Sork et al. 1993, Kelly 1994, Kelly and Sork 2002), has been shown to influence the population dynamics of seed predators and associated species (e.g., Ostfeld et al. 1996, McShea 2000) and may influence the risk of post-dispersal predation to seeds and fruits (Silvertown 1980).

MATERIALS AND METHODSChemical data to test the non-profitable resource hypothesis were sum-

marized from a range of studies (Table 2) and focused on four main nut defense/nutritional parameters (concentrations of tannin, crude fat, crude protein, and crude carbohydrate) expressed on a percent dry weight basis. Chemical data were summarized for eight nut-bearing tree species in each of three taxonomic groups (white oaks, subgenus Lepidobalanus; red oaks, subgenus Erythrobal-anus; and hickories, Juglandaceae) that occur within the range of American beech and for which adequate data were available. Using data from individual species, means were calculated for each chemical parameter by nut tree taxo-nomic group. When multiple values for a chemical parameter were available for a tree species, they were averaged and the species mean was used in calculating the taxonomic group mean. In instances where the concentration of a chemical constituent was listed as trace or negligible, a default value of 0.1% was used in the analysis.

I used both univariate and multivariate statistical approaches to analyze nut defense/nutritional data. Univariate tests allowed me to explore potential differences in single nut defense/nutritional parameters between American beech and the three taxonomic groups of nut trees as described above. Uni-variate one sample t-tests (α ≤ 0.05) were used to explore differences in nut chemistry between American beech and each of the three taxonomic groups of trees. Multivariate principal components analysis (PCA) was used to examine the relationship of the suite of nut nutritional and chemical defense parameters across tree species and not just single factors as in univariate tests. One sample t-tests were done using SYSTAT version 7.0 (Wilkinson 1997). PCA was done using MVSP version 3.14 (Kovach 2000).

Data to test the unpredictable resource hypothesis were obtained from a ten-year American beech mast crop study conducted by Gysel (1971) in southern


ble

1. S

peci

es ri

chne

ss o

f Nor

th A

mer

ican

nut

-infe

stin

g in

sect

spe

cies

by

plan

t fam

ily a

nd g

enus

(com

pile

d fr

om W

illia

ms

1989

). C

asta

no-

psis

and

Lith

ocar

pus a

re m

uch

mor

e re

stri

cted

in ra

nge

than

the

othe

r nut

tree

gen

era

(bot

h ar

e co

nfine

d to

a n

arro

w co

asta

l ban

d of

Cal

ifor-

nia,

Ore

gon

and

Was

hing

ton,

USA

) but

are

incl

uded

for c

ompa

rativ

e pu

rpos

es.

Num

ber

of s

peci

es b

y in

sect

ord

er

N

umbe

r of

spe

cies

by

host

taxo

n

Hos

t pla

nt fa

mily

/gen

us

Col

eopt

era

Dip

tera

Le

pido

pter

a H

ymen

opte

ra

Gen

us

Fam

ily

Betu

lace

ae

Cor

ylus

L.

2 0

1 0

3 3

Faga

ceae

C

asta

nea

Mill

. 5

0 2

0 7

76

Cas

tano

psis

(D.D

on) S

prac

h 1

0 1

0 2

Fa

gus

L.

0 0

1 0

1

Lith

ocar

pus B

lum

e 1

0 1

0 2

Q

uerc

us L

. 24

2

4 34

64

Jugl

anda

ceae

C

arya

Nut

t. 3

1 3

0 7

16

Jugl

ans L

. 2

4 3

0 9

Tota

l by

inse

ct o

rder

38

7

16

34

95


ble

2. C

hem

ical

and

mas

t fre

quen

cy d

ata

for

sele

cted

nut

s of

Nor

th A

mer

ican

nut

-bea

ring

tree

s ex

pres

sed

on a

per

cent

dry

wei

ght b

asis

. Su

pers

crip

ts re

fer t

o re

fere

nces

from

whi

ch d

ata

wer

e ob

tain

eda (

rang

e of

val

ues r

ecor

ded

acro

ss st

udie

s app

ear i

n pa

rent

hese

s). D

ata

on m

ast

crop

freq

uenc

y (m

ast)

is fr

om S

chop

mey

er (1

974)

and

is p

rese

nted

as a

rang

e. F

or ta

nnin

conc

entr

atio

n in

Fag

us g

rand

ifolia

and

Jug

land

acea

e,

a da

shed

line

indi

cate

s th

at a

neg

ligib

le a

mou

nt (<

0.5

%) w

as d

etec

ted.

NA

= da

ta n

ot a

vaila

ble;

crud

e ca

rb. =

crud

e ca

rboh

ydra

te.

Taxo

n Ta

nnin

(%

) C

rude

fat (

%)

Cru

de p

rote

in (%

) C

rude

car

b. (%

) M

ast (

year

s)

Fagu

s gr

andi

folia

Ehr

h.

---

10.6

5 7.

85 6.

55 2-

8

Faga

ceae

: Lep

idob

alan

us

Que

rcus

alb

a L.

4.

5 (3

.3-5

.6)1,

13

5.7

(2.9

-8.8

)1,3,

7,8,

13

5.7

(4.6

-7.3

)1,3,

8,13

46

.63

4-10

Q. b

icol

or W

illd.

2.

16 2.

98 4.

46,8

NA

3-5

Q. l

yrat

a W

alt.

0.66

0.98

4.5

(4.4

-4.6

)6,8

49.8

5 3-

4Q

. mac

roca

rpa

Mic

hx.

2.0

(0.7

-3.2

)6,12

6.

7 (4

.8-9

.8)5,

7,13

4.

4 (3

.9-4

.9)5,

6,12

45

.95

2-3

Q. m

uehl

enbe

rgii

Enge

lm.

4.712

6.

4 (6

.1-6

.6)3,

12

4.6

(4.4

-4.8

)3,12

34

.53

NA

Q. p

rino

ides

Will

d.

4.41

6.31

7.61

NA

NA

Q. p

rinu

s L.

9.3

(8.1

-10.

4)1,

13

6.8

(3.3

-10.

1)1,

2,10

,13

6.3

(5.8

-6.9

)1,2,

10,1

3 N

A 2-

3Q

. ste

llata

Wan

genh

. 3.

7 (0

.9-6

.5)6,

12

5.9

(5.2

-6.7

)2,3,

8,12

5.

5 (3

.8-6

.8)2,

3,6,

8,12

37

.93

2-3

Mea

n ±

1 SE

3.

9 ±

0.9

5.2

± 0.

8 5.

4 ±

0.4

42.9

± 2

.9

Faga

ceae

: Ery

thro

bala

nus

Que

rcus

falc

ata

Mic

hx.

8.76

18.4

(15.

6-22

.7)2,

5,8

5.8

(4.2

-7.0

)2,5,

6,8

40.4

(23.

0-57

.7)2,

5 1-

2Q

. ili

cifo

lia W

ange

nh.

1.31

19.7

(19.

4-20

.0)1,

2

8.2

(6.1

-10.

3)1,

2

NA

NA

Q. m

arila

ndic

a M

uenc

hh.

76 10

.72

6.6

(6.3

-6.9

)2, 6

N

A N

AQ

. nig

ra L

. 8.

86 20

.7 (2

0.3-

21.1

)3,8

4.7

(3.8

-5.4

)3,6,

8 25

.83

1-2

Q. p

alus

tris

Mue

nchh

. 9.

312

13.6

(11.

7-15

.4)5,

12

5.0

(3.8

-6.1

)5, 1

2 45

.45

1-2

Q. p

hello

s L.

7.26

19.8

(19.

6-20

.0)3,

8 5.

5 (5

.2-5

.9) 3

,6,8

31

.23

1Q

. rub

ra L

. 11

.5 (9

.8-1

3.0)

1,10

,12

19.0

(14.

0-23

.0)1,

8, 1

2,13

5.

9 (4

.9-6

.6)1,

8, 1

2,13

N

A 3-

5Q

. vel

utin

a La

m.

16.5

12

15.3

(13.

0-17

.5)8,

12

5.9

(5.7

-6.0

) 8,1

2 N

A 2-

3

Mea

n ±

1 SE

9.

9 ±

1.2

17.2

± 1

.3

6.0

± 0.

4 34

.1 ±

5.9


ble

2. C

ontin

ued.

Taxo

n Ta

nnin

(%

) C

rude

fat (

%)

Cru

de p

rote

in (%

) C

rude

car

b. (%

) M

ast (

year

s)

Jugl

anda

ceae

C

arya

aqu

atic

a (M

ichx

. F.)

Nut

t. ---

32

.09

10.2

9 54

.29

1-2

C. c

ordi

form

is

(Wan

genh

.) K

. Koc

h ---

39

.6 (3

0.8-

48.3

)5,9

5.4

(3.3

-7.5

)5,9

29.2

(17.

1-41

.3)5,

9 3-

5C

. flor

idan

a Sa

rg.

---

34.3

11

9.611

45

.311

N

AC

. illi

noen

sis

(W

ange

nh.)

K. K

och

---

32.9

3 9.

33 13

.33

1-2

C. l

acin

iosa

(M

ichx

. F.)

Loud

on

---

8.73

1.53

13.8

3 1-

2C

. myr

istic

iform

is

(Mic

hx. F

.) N

utt

.---

15.2

3 5.

83 16

.13

2-3

C. o

vata

(Mill

.) K

. Koc

h 0.

51 33

.4 (2

9.3-

37.4

)3,7,

9 10

.8 (5

.9-1

3.3)

1,3,

9 10

.9 (8

.8-1

3.0)

1,3

1-

3C

. tom

ento

sa (P

oir.)

Nut

t .--

- 20

.05

3.75

12.7

5 2-

3

Mea

n ±

1 SE

---

27

.0 ±

3.9

7.

0 ±

1.2

24.4

± 5

.9

a Ref

eren

ces

incl

ude:

1 Wai

no a

nd F

orbe

s (1

941)

, 2 Kin

g an

d M

cClu

re (1

944)

, 3 Bon

ner (

1971

), 4 B

urns

and

Vie

rs (1

973)

, 5 Bon

ner (

1974

), 6 O

fcar

cik

and

Burn

s (1

971)

, 7 Sm

ith a

nd F

ollm

er (1

972)

, 8 Sho

rt (1

976)

, 9 Hal

ls (1

977)

, 10Sm

allw

ood

and

Pete

rs (1

986)

, 11Ab

raha

mso

n an

d Ab

raha

mso

n (1

989)

, 12Br

iggs

and

Sm

ith (1

989)

, 13Se

rvel

lo a

nd K

irkp

atri

ck (1

989)

.


Michigan, USA, and a six-year study conducted by Leak (1993) in the White Mountains of New Hampshire, USA. I lumped Gysel’s (1971) and Leak’s (1993) nut condition classes - sound (i.e., non-infested or damaged), insect-infested, and vertebrate-damaged - into a single class, available mast, for analysis. Neither Gysel (1971) nor Leak (1993) identified the specific insect species infesting beechnuts at their study site. However, their descriptions of frass-filled nuts, characteristic of nut-infesting lepidopterans, strongly implicate C. latiferreana (e.g., Winston 1956, Gibson 1971). Moreover, Graber and Leak (1992), in a related study in New Hampshire, identified C. latiferreana as the sole insect predator of beech nuts. Pearson product-moment correlation (α ≤ 0.05) with Bon-ferroni correction was used to examine the relationships of insect-infestation and vertebrate damage to beechnuts with available mast across years. Percentage data were arcsin transformed prior to analysis to ensure normality (Zar 1996). Correlation analysis was done using SYSTAT version 7.0 (Wilkinson 1997).

RESULTSResults from univariate statistical tests suggest that nuts of American

beech were more similar in tannin concentration and nutritional value to nuts of the hickory group than to those of either the white oak or red oak groups (Tables 2 and 3). Beechnuts were significantly lower in crude fat and crude carbohydrate concentrations than hickory nuts but did not differ significantly in tannin or crude protein concentrations (Table 3). Beechnuts had significantly higher crude fat and protein concentrations than acorns of the white oak group but were significantly lower in tannin and crude carbohydrate concentrations (Table 3). Beechnuts had a significantly higher concentration of crude protein than red oak acorns but were significantly lower in tannin, crude fat, and crude carbohydrate concentrations (Table 3). Overall, beechnuts were consistently lower in crude carbohydrate concentration, generally lower in crude fat and tannin concentrations, and generally higher in crude protein concentration, than nuts of the white oak, red oak, and hickory species groups examined in this study.

PCA showed a clear separation of most hickory species from white and red oak species largely on the basis of crude fat, crude protein and tannin con-tent of nuts (Fig. 1). Red and white oak species groups were separated from each other in ordination space mostly on the basis of high tannin content (red oaks) and high carbohydrate content (white oaks). Principal components axis 2 accounted for 41.6% of the variance in the data matrix and separated nuts of species on a gradient from high tannin concentration to high crude fat and high crude protein percentages (Table 4). Principal components axis 1 accounted for 30.7% of the variance in the data matrix and separated nuts of species on a gradient from high tannin to high crude carbohydrate percentages. Together the two principal components axes accounted for 72.3% of the variance in the data matrix. American beech clustered in the middle of ordination space (Fig. 1) suggesting that it is intermediate in the suite of the four nut nutrition and defense parameters considered in this study.

American beech exhibited the second greatest variation in frequency of mast production of any of the nut tree species examined (Table 2). Gysel (1971) observed a large mast crop of viable beechnuts only once in ten years and crop failures twice (Fig. 2). Sound nuts comprised less than 10% of the total beechnut crop for 7 of 10 years (mean = 15.2 ± 5.0% SE, range = 2.4 to 47.5%). Incomplete, nonviable nuts comprised more than 20% of the annual production for nine years (mean = 43.5 ± 7.6% SE, range = 23.5 to 87.7%). Yearly variance in nut crops (65%), and variance among individual trees (30%), accounted for most of the variation in beechnut crop production during Gysel’s (1971) study. In contrast, Leak (1993) found that sound nuts comprised an average of 80% (± 2.5 SE) of the annual production during his six-year-study with no nut crop failures.


ble

3. C

ompa

riso

n of

chem

ical

cons

titue

nts

of n

uts

of A

mer

ican

bee

ch (A

B) w

ith th

ose

of th

e w

hite

oak

(WO

), re

d oa

k (R

O),

and

hick

ory

(HK

) spe

cies

gro

ups.

Sig

nific

ant d

iffer

ence

s (P

< 0

.05,

one

sam

ple

t-tes

t) be

twee

n gr

oups

are

den

oted

by

< an

d >;

non

-sig

nific

ance

is d

enot

ed

by =

.

Tann

in (%

) C

rude

fat (

%)

Cru

de p

rote

in (%

) C

rude

car

bohy

drat

e (%

)

AB <

WO

AB

> W

O

AB >

WO

AB

< W

O

(P =

0.0

05, d

f =7,

t =

4.11

) (P

< 0

.000

1, d

f =7,

t =

-7.1

5)

(P =

0.0

01, d

f =7,

t =

-5.9

9)

(P =

0.0

05, d

f =7,

t =

12.6

5)

AB <

RO

AB

< R

O

AB >

RO

AB

< R

O

(P <

0.0

001,

df =

7, t

= 8.

39)

(P =

0.0

01, d

f =7,

t =

5.20

) (P

= 0

.002

, df =

7, t

= -4

.84)

(P

= 0

.007

, df =

3, t

= 6.

61)

AB =

HK

AB

< H

K

AB =

HK

AB

< H

K

(P =

0.5

, df =

7, t

= 0.

73)

(P =

0.0

04, d

f =7,

t =

4.25

) (P

= 0

.6, d

f =7,

t =

-0.6

3)

(P =

0.0

19, d

f =7,

t =

3.02

)


Insect damage to beechnut crops in Gysel’s (1971) study ranged from 1.2 to 23.3% (mean = 10.7 ± 2.5% SE). Vertebrate damage to beechnut crops was greater than insect damage, and ranged from 0.6 (crop failure year) to 46.2% of the total crop (mean = 27.8 ± 5.6% SE). Both insect infestation (r = 0.82, df = 1, P = 0.004) and vertebrate damage (r = 0.94, df = 1, P < 0.0001) were sig-nificantly and positively correlated with available beechnut mast across study years. However, during the two peak years of sound beechnut production, insect damage was low (3.9% and 7.6% of the total crop; Figure 2).

Insect damage to beechnut crops in Leak’s (1993) study ranged from 16.0 to 100% (mean = 62.0 ± 12.0 % SE). Vertebrate damage to beechnut crops was less than insect damage, and ranged from 0 (an apparent outbreak year in which insects destroyed the whole nut crop) to 20.0% of the total crop (mean = 8.5 ± 2.7% SE). Both insect infestation (r = 0.85, df = 1, P = 0.034) and vertebrate damage (r = 0.85, df = 1, P < 0.031) were significantly and positively correlated with available beechnut mast across study years.

DISCUSSIONI found no strong evidence to suggest that chemical defense or low nutri-

tional value was associated with the species-poor nut-infesting insect fauna of American beech. In contrast, beechnuts appear to be a quality food resource for nut-feeding insects, having good protein content and negligible levels of tannins.

Masting, a complicated phenomenon influenced by weather, past reproduc-tive events and root carbohydrate reserves (Matthews 1955, Sork et al. 1993, Piovesan and Adams 2001, Kelly and Sork 2002), is widespread in the Fagaceae, particularly among oaks (e.g., Downs and McQuilken 1944, McShea 2000, Table 2). As in American beech, nut crop failure is not uncommon in oaks (Downs and McQuilken 1944, Sork et al. 1993, McShea 2000), thus year-to-year variability in nut crop size alone does not explain the great difference in species richness of the nut-infesting insect fauna between these two taxa. In contrast, American beech and hickories, somewhat similar in nut tannin content and nutritional value, differ greatly in mast periodicity (Nixon et al. 1980, Sork 1983; Table 1). Hickory nuts, produced at relatively frequent intervals across years, may be a more predictable, easily colonized food resource for insects than are nutrition-ally comparable but temporally variable beechnuts.

I suggest that the great variability in production of sound versus incom-plete beechnuts both within crops and across years as noted by Gysel (1971; Fig. 1) and others (Ward 1961, Dix and Skrentny, Jr. 1965, Stalter 1982, Johnson and Adkisson 1985) has restricted colonization of nuts of American beech by insects and limited accrual of species. An incomplete beechnut lacks endosperm and

Table 4. Results of principal components analysis (PCA) for nut nutritional and chemi-cal defense parameters.

PCA variable loadings Axis 1 Axis 2

% Tannin content -0.360 -0.574 % Crude fat 0.666 -0.183 % Crude protein 0.625 -0.354 % Crude carbohydrates 0.191 0.716Eigenvalues 1.663 1.229Percent variance explained 41.57 30.72

148 THE GREAT LAKES ENTOMOLOGIST Vol. 40, Nos. 3 & 40.

6

0.5

0.4

0.2

0.1

-0.1

-0.2

-0.4

-0.5

-0.6

-0.6

-0

.5

-0.4

-0

.2

-0.1

0.

1 0.

2 0.

4 0.

5 0.

6

Axi

s 1

Axis 2

TANNIN

CRUCAR

CRUFAT

CRUPRO

CARC

ORCA

RAQU

CARF

LO

CARO

VA

CARI

LL

QUEI

LIQU

EVEL

QUEP

RN

QUER

UB

QUEN

IG

QUEB

IC

CARL

AC

FAGG

RA

QUEM

ARQU

EPRI

QUEA

LBQU

EMAC

QUEL

YR

QUEM

ULQU

ESTE

QUEF

AL QUEP

PE

CART

OMCA

RMYR

QUEP

AL

Vect

or s

calin

g 0.

86

Figu

re 1

. Pr

inci

pal c

ompo

nent

s bi

plot

for n

ut d

efen

se/n

utri

tiona

l cha

ract

ers

by tr

ee s

peci

es.

Spec

ies

are

from

Tab

le 2

and

are

code

d by

the

first

thre

e le

tter

s of

the

genu

s an

d th

e fir

st th

ree

lett

ers

of th

e sp

ecifi

c epi

thet

. Li

nes

are

vect

ors

of n

ut d

efen

se/n

utri

tiona

l par

amet

ers

asso

-ci

ated

with

pri

ncip

al c

ompo

nent

s ax

es.

CRU

CAR

= cr

ude

carb

ohyd

rate

conc

entr

atio

n, C

RUFA

T =

crud

e fa

t con

cent

ratio

n, C

RUPR

O =

crud

e pr

otei

n co

ncen

trat

ion,

TAN

NIN

= ta

nnin

conc

entr

atio

n.


an embryo, but has a fully developed pericarp and is morphologically similar to sound nuts (Gysel 1971). Incomplete nuts are also produced by European beech (Matthews 1955, Hilton and Packham 1986). Oaks also produce incom-plete or abortive nuts, but these generally comprise less of the total crop than incomplete beechnuts and are small and deformed in appearance compared to sound nuts (Downs and McQuilken 1944, Sork et al. 1983). Vertebrates, such as blue jays, Cyanocitta cristata (L.), can discriminate between sound and incomplete beechnuts and preferentially select sound nuts for feeding and caching (Johnson and Adkisson 1985), presumably by tactile and visual means as in other corvids (e.g., Ligon and Martin 1974). However unlike vertebrates, nut-infesting insects may be limited in their ability to find sound beechnuts in a large crop of incomplete nuts or they may be unable to distin-guish between sound and incomplete nuts (e.g., Hall et al. 1979, Butkewich et al. 1987, Desouhant 1998, Stamps and Linit 2002). Adult nut insects may select incomplete beechnuts that cannot provide the energy needed for larval growth and development or alternatively, they may expose themselves to in-creased predation risk when searching tree canopies for spatially dispersed, numerically rare, sound beechnuts.

The life history of C. latiferreana provides further evidence that variabil-ity in beechnut crops may have restricted development of a diverse nut insect fauna on American beech. Compared to most primary nut insects (i.e., those capable of entering nuts through their own feeding or oviposition holes, Win-ston 1956), C. latiferreana feeds on a broad range of hosts, including nuts of 19 tree and shrub species in 6 genera and 3 families as well as the fleshy fruits of other woody plants (Dohanian 1940, Williams 1989). Other insect species, like Conotrachelus and Curculio acorn weevils, are nut specialists whose hosts are typically confined to 1 or 2 genera in a single family (Williams 1989). A broad host range is a means by which C. latiferreana can cope with unpredictable beechnut resources by switching to alternate, more abundant food resources

Figure 2. Mean annual production of sound, incomplete, vertebrate-damaged, and insect-infested nuts of American beech per 100 ft2 (9.3 m2) of crown collection area in Michigan, USA. Data are from a ten-year mast crop study conducted by Gysel (1971). Data were collected from 20 trees in 2 woodlots for 2 years and 30 trees in three wood-lots for 8 years.


when necessary. Moreover, larvae of C. latiferreana, unlike those of most other nut-feeding insects, have some mobility and can relocate to different nuts when faced with diminished food resources (Winston 1956, Gibson 1971).

It is interesting to speculate why other North American nut-infesting insect species besides C. latiferreana apparently failed to develop generalist feeding strategies under the selective pressure of masting. Perhaps phylo-genetic constraints in the plasticity of certain morphological, physiological or behavioral traits limited the development of generalist feeding strategies in the other nut-infesting taxa. A second possibility is that C. latiferreana may be a superior competitor that eliminated other nut-feeding species through competi-tive exclusion. Finally, perhaps C. latiferreana had a limited pool of natural enemies to limit is population size and allow for coexistence of other generalist nut-feeding insect species.

It is also important to consider the competitive effects of other nut-consuming animals on beechnut resources and their potential influence on the accrual of a diverse nut insect fauna. Beechnuts are widely used as food by many species of North American vertebrates including several species of tree squirrels (Sciurus, Tamiasciurus), blue jay, C.cristata , wild turkey, Meleagris gallopavo (L.), ruffed grouse, Bonasa umbellatus (L.), white-tailed deer (Odo-coileus virginianus Zimmermann), black bear (Ursus americanus Pallas), and other birds and mammals (Martin et al. 1951, Nixon et al. 1968, Halls 1977, Johnson and Adkisson 1985, Webb 1986). Gysel (1971) and others (Graber and Leak 1992, Leak 1993) have noted that sound beechnuts fallen beneath trees were quickly consumed or removed by vertebrates and very few nuts survived more than two to three weeks. Likewise, harvest of beechnuts from beech canopies by blue jays can be extensive (Johnson and Adkisson 1985). Selective harvest by vertebrates would further reduce the number of sound beechnuts available to insects and increase the probability that insects within nuts themselves may fall prey to vertebrates. Vertebrate nut predators like the white-footed mouse (Peromyscus leucopus Rafinesque) and grey squirrel (Sciurus carolinensis) generally cannot discriminate between non-infested and insect-infested nuts and will feed on either (Semel and Andersen 1988, Weckerly et al. 1989). It should be noted that contemporary nut harvests by vertebrates are nowhere near the magnitude of historic harvests by massive flocks of the extinct passenger pigeon, Ectopistes migratorius (L.), for which beechnuts were a preferred food (Schorger 1955, Webb 1986, Ellsworth and McComb 2003). Whether competition for beechnut resources with passenger pigeons influenced the composition of the present-day nut insect fauna of American beech can only be speculated.

Based on the arguments outlined above, I suggest that spatial and temporal unpredictability of the nut crop of American beech was an effective filter limiting colonization of beechnuts by insects and the accrual of a diverse insect fauna. The lone insect species able to successfully colonize beechnuts, C. latiferreana, has a relatively broad host range that buffers it from resource unpredictability, unlike specialist insect species that infest nuts of few hosts. Given the parallels between European beech and American beech in mast frequency, production of incomplete nuts, and a species-poor nut insect fauna dominated by Cydia species (Matthews 1955, Nielsen 1977, Jensen 1985, Nils-son 1985, Hilton and Packham 1986, Piovesan and Adams 2001), I also suggest that unpredictability of beechnut resources helped to shape the composition of the nut insect fauna of European beech.

ACKNOWLEDGMENTSThe helpful comments of two anonymous reviewers improved the content

of this paper.


LITERATURE CITEDAbrahamson, W. G., and C. R. Abrahamson. 1989. Nutritional quality of animal dispersed

fruits in Florida sandhill ridges. Bull. Torrey Bot. Club 116: 215-228.Andersen, A. N., and T. R. New. 1987. Insect inhabitants of fruits of Leptospermum,

Eucalyptus and Casuarina in south-eastern Australia. Aust. J. Zool. 35: 327-336.Blaustein, A. R., A. M. Kuris, and J. J. Alio. 1983. Pest and parasite species-richness

problems. Am. Nat. 122: 556-566.Bonner, F. T. 1971. Chemical contents of southern hardwood fruits and seeds. USDA

Forest Service Res. Note SO-136.Bonner, F. T. 1974. Chemical components of some southern fruits and seeds. USDA

Forest Service Res. Note SO-183.Briggs, J. M., and K. G. Smith. 1989. Influence of habitat on acorn selection by Peromy-

scus leucopus. J. Mamm. 70: 35-43.Burns, T. A., and C. E. Viers. 1973. Caloric and moisture content values of selected fruits

and mast. J. Wildl. Manage. 37: 585-587.Butkewich, S. L., R. J. Prokopy, and T. A. Green. 1987. Discrimination of occupied

host fruit by plum curculio females (Coleoptera: Curculionidae). J. Chem. Ecol. 13: 1833-1841.

Desouhant, E. 1998. Selection of fruits for oviposition by the chestnut weevil, Curculio elephas. Entomol. Exp. Appl. 86:71-78.

Dix, R. L., and R. F. Skrentny, Jr. 1965. Reproduction of Fagus grandifolia Ehrh. by seed in Wisconsin. Can. J. Bot. 43: 757-763.

Dohanian, S. M. 1940. Melissopus latiferreanus as a pest of filberts in the northwest. J. Econ. Entomol. 33:852-856.

Downs, A. A., and W. E. McQuilken. 1944. Seed production of southern Appalachian oaks. J. For. 42: 913-920.

Ellsworth, J. W., and B. C. McComb. 2003. Potential effects of passenger pigeon flocks on the structure and composition of presettlement forests of eastern North America. Conserv. Biol. 17: 1548-1558.

Gibson, L. P. 1971. Insects of bur oak acorns. Ann. Entomol. Soc. Am. 64:232-234.Gleason, H. A., and A. Cronquist. 1991. Manual of vascular plants of northeastern

United States and adjacent Canada. 2nd edition. The New York Botanical Garden, Bronx, NY.

Graber, R. E., and W. B. Leak. 1992. Seed fall in an old-growth northern hardwood forest. USDA For. Serv., Northeastern For. Exp. Sta., Res. Pap. NE-663.

Gysel, L. W. 1971. A 10-year analysis of beechnut production and use in Michigan. J. Wildl. Manage. 35:516-519.

Hall, M. J., R. D. Eikenbary, and W. D. Warde. 1979. Effects of pecan cluster size on the selection of nuts for feeding and oviposition by the pecan weevil, Curculio caryae (Coleoptera: Curculionidae). Can. Entomol. 111: 1193-1196.

Halls, L. K. 1977. Southern fruit-producing woody plants used by wildlife. USDA For. Serv. Gen. Tech. Rep. SO-16.

Hilton, G. M. and J. R. Packham. 1986. Annual and regional variation in English beech mast (Fagus sylvatica L.). Arboric. J. 10: 3-14.

Jensen, T. S. 1985. Seed-seed predator interactions of European beech, Fagus silvatica and forest rodents, Clethrionomys glareolus and Apodemus flavicollis. Oikos 44: 149-156.

Johnson, W. C., and C. S. Adkisson. 1985. Dispersal of beech nuts by blue jays in frag-mented landscapes. Am. Midl. Nat.113: 319-324.


Kelly, D. 1994. The evolutionary ecology of mast seeding. Trends Ecol. Evol. 9: 465-470.Kelly, D., and V. L. Sork. 2002. Mast seeding in perennial plants: what, how, where?

Ann. Rev. Ecol. Syst. 33: 427-447.King, T. R., and H. E. McClure. 1944. Chemical composition of some American wild

feedstuffs. J. Agric. Res. 69: 33-46.Kovach, W. L. 2000. MVSP: A Multivariate Statistical Package for Windows. Version

3.14. Kovach Computing Services, Pentraeth, Wales, UK.Leak, W. B. 1993. Six-year beechnut production in New Hampshire. USDA For. Serv.

Northeastern For. Exp. Stat. Res. Pap. NE-677.Ligon, J. D., and D. J. Martin. 1974. Pinon seed assessment by the pinon jay, Gymnorhi-

nus cyanocephalus. Anim. Behav. 22: 421-429.Marquis, D. A., P. L. Eckert, and B. A. Roach. 1976. Acorn weevils, rodents, and deer

all contribute to oak-regeneration difficulties in Pennsylvania. USDA For. Serv. Res. Pap. NE-356.

Martin, A. C., H. S. Zim, and A. L. Nelson. 1951. American wildlife and plants: a guide to wildlife food habits. Dover Publications., Inc., New York.

Martin, J. S., and M. M. Martin. 1982. Tannin assays in ecological studies: lack of cor-relation between phenolics, proanthocyanidins and protein-precipitating constituents in mature foliage of six oak species. Oecologia 54: 205-211.

Matthews, J. D. 1955. The influence of weather on the frequency of beech mast years in England. Forestry 28: 107-116.

McCoy, E. D., and J. R. Rey. 1983. The biogeography of herbivorous arthropods: species accrual on tropical crops. Ecol. Entomol. 8: 305-313.

McShea, W. J. 2000. The influence of acorn crops on annual variation in rodent and bird populations. Ecology 81: 228-238.

Nielsen, B. O. 1977. Beech seeds as ecosystem components. Oikos 29: 268-274.Nilsson, S. G. 1985. Ecological and evolutionary interactions between reproduction of

beech Fagus silvatica and seed eating animals. Oikos 44: 157-164.Nixon, C. M., D. M. Worley, and M. W. McClain. 1968. Food habits of squirrels in south-

east Ohio. J. Wildl. Manage. 32: 294-305.Nixon, C. M., M. W. McClain, and L. P. Hansen. 1980. Six years of hickory seed yields

in southeastern Ohio. J. Wildl. Manage. 44: 534-539.Ofcarcik, R. P., and E. E. Burns. 1971. Chemical and physical properties of selected

acorns. J. Food Sci. 36: 576-578.Ostfeld R. S., C. G. Jones, and J. O. Wolff. 1996. Of mice and mast: ecological connections

in eastern deciduous forests. BioScience 46: 323-330.Piovesan G., and J. J. Adams. 2001. Masting behaviour in beech: linking reproduction

and climatic variation. Can. J. Bot. 79: 1039-1047.Schopmeyer, C. S. (tech. coord.) 1974. Seeds of woody plants in the United States. USDA

Agriculture Handbook No. 450, Washington, DC.Schorger, A. W. 1955. The passenger pigeon: its natural history and extinction. University

of Wisconsin Press, Madison, WI.Semel, B., and D. C. Andersen. 1988. Vulnerability of acorn weevils (Coleoptera: Curcu-

lionidae) and attractiveness of weevils and infested Quercus alba acorns to Peromyscus leucopus and Blarina brevicauda. Am. Midl. Nat. 119: 385-393.

Servello, F. A., and R. L. Kirkpatrick. 1989. Nutritional value of acorns for ruffed grouse. J. Wildl. Manage. 53: 26-29.

Short, H. L. 1976. Composition and squirrel use of acorns of black and white oak groups. J. Wildl. Manage. 40: 479-483.


Silvertown, J. W. 1980. The evolutionary ecology of mast seeding in trees. Biol. J. Linn. Soc. 14: 235-250.

Smallwood, P. D., and W. D. and Peters. 1986. Grey squirrel food preferences: the effects of tannin and fat concentrations. Ecology 67: 168-174.

Smith, C. C., and D. Follmer. 1972. Food preferences of squirrels. Ecology 53: 82-91.Sork, V. L. 1983. Mast-fruiting in hickories and availability of nuts. Am. Midl. Nat.

109: 81-88.Sork, V. L., P. Stacey, and J. E. Averett. 1983. Utilization of red oak acorns in non-bumper

crop year. Oecologia 59: 49-53.Sork, V. L., J. Bramble, and O. Sexton. 1993. Ecology of mast-fruiting in three species

of North American deciduous trees. Ecology 74: 528-541.Stalter, R. 1982. Production of viable seed by the American beech (Fagus grandifolia).

Bull. Torrey Bot. Soc. 109: 542-544.Stamps, W. T., and M. J. Linit. 2002. Oviposition choice by the black walnut curculio,

Conotrachelus retentus (Coleoptera: Curculionidae): a ten year study. Environ. Entomol. 32: 281-284.

Strong, D. R., Jr. 1974. Rapid asymptotic species accumulation in phytophagous insect communities: the pests of cacao. Science 185: 1064-1066.

Strong, D. R., Jr. 1979. Biogeographic dynamics of insect-host plant communities. Ann. Rev. Ecol. Syst. 24: 89-119.

Strong, D. R., Jr., J. H. Lawton, and T. R. E. Southwood. 1984. Insects on plants: com-munity patterns and mechanisms. Harvard University Press, Cambridge, MA.

Wainio, W. W., and E. B. Forbes. 1941. The chemical composition of forest fruits and nuts from Pa. J. Agric. Res. 62: 627-635.

Ward, R. T. 1961. Some aspects of the regeneration habits of the American beech. Ecol-ogy 42: 828-832.

Watt, A. S. 1923. On the ecology of British beechwoods with special reference to their regeneration. J. Ecol. 11: 1-48.

Webb, S. L. 1986. Potential role of passenger pigeons and other vertebrates in the rapid Holocene migrations of nut trees. Quaternary Res. 26: 367-375.

Weckerly, F., K. E. Nicholsen, and R. D. Semlitsch. 1989. Experimental test of discrimi-nation by squirrels for insect-infested and noninfested acorns. Am. Midl. Nat. 122: 412-415.

Wilkinson, L. 1997. SYSTAT 7.0. Statistics. SPSS, Inc., Chicago, IL.Williams, C. E. 1989. Checklist of North American nut-infesting insects and host plants.

J. Entomol. Sci. 24: 550-562.Winston, P. W. 1956. The acorn microsere, with special reference to arthropods. Ecology

37: 120-132.Zar, J. H. 1996. Biostatistical analysis. 3rd ed. Prentice-Hall, Inc., Upper Saddle River,

NJ.


1Department of Biological Sciences, Western Illinois University, 1 University Circle, Macomb, IL 61455. 2U.S. Geological Survey, 160 N. Stephanie St., Henderson, NV 89074.

EFFECTS OF PITFALL TRAP PRESERVATIVE ON COLLECTIONS OF CARABID BEETLES (COLEOPTERA: CARABIDAE)

Kenneth W. McCravy1 and Jason E. Willand2

ABSTRACTEffects of six pitfall trap preservatives (5% acetic acid solution, distilled

water, 70% ethanol, 50% ethylene glycol solution, 50% propylene glycol solution, and 10% saline solution) on collections of carabid beetles (Coleoptera: Carabidae) were studied in a west-central Illinois deciduous forest from May to October 2005. A total of 819 carabids, representing 33 species and 19 genera, were collected. Saline produced significantly fewer captures than did acetic acid, ethanol, eth-ylene glycol, and propylene glycol, while distilled water produced significantly fewer captures than did acetic acid. Significant associations between numbers of captures and treatment were seen in four species: Amphasia interstitialis (Say), Calathus opaculus LeConte, Chlaenius nemoralis Say, and Cyclotrachelus sodalis (LeConte). Results of this study suggest that type of preservative used can have substantial effects on abundance and species composition of carabids collected in pitfall traps.

____________________

Pitfall trapping is a commonly used method of sampling surface-active soil and litter arthropods such as carabid beetles (Greenslade 1964, Holopainen 1992, Lemieux and Lindgren 1999), rove beetles, Staphylinidae (Honêk 1988), ants, Formicidae (Greenslade 1973), and wandering spiders such as wolf spi-ders, Lycosidae (Curtis 1980, Honêk 1988). Pitfall trap collections reflect an interaction between arthropod activity and abundance (Thiele 1977), however, there is evidence that different arthropod species perceive and respond to pitfall traps differently and that trap characteristics can affect capture rates (Halsall and Wratten 1988, Digweed et al. 1995, Work et al. 2002). Pitfall traps are often used with a preservative/killing agent to maintain the condition of the trapped specimens and to reduce escape and within-trap predation. One factor that can affect arthropod response to pitfall traps is the type of preservative used. A wide variety of preservatives have been used in pitfall traps, includ-ing ethylene glycol, propylene glycol, water, formalin, kerosene, brine, alcohol, acetic acid, chloral hydrate, and benzoic/acetic acid (Woodcock 2005); however, the type of preservative used can affect the number, species, or even sex ratio of arthropod captures. For carabids, effects on pitfall trap collections have been found for ethylene glycol (Holopainen 1990, 1992), propylene glycol (Hammond 1990), benzoic/acetic acid (Scheller 1984), and formalin (Luff 1968, Scheller 1984; Holopainen and Varis 1986), although Waage (1985) found no evidence of formalin influencing collections.

Carabidae is one of the most diverse insect families, with over 40,000 described species (Lövei and Sunderland 1996). Carabids are important preda-tors in many terrestrial ecosystems, and can be important biological control agents (Lövei and Sunderland 1996). The ecology and behavior of carabids are often closely associated with factors such as soil type, vegetation cover and microclimate, making them potentially important bioindicators (Thiele 1977,


Niemelä et al. 1992, Ings and Hartley 1999, Villa-Castillo and Wagner 2002, McCravy and Willand 2005, Willand and McCravy 2006). Knowledge of the potential effects of type of preservative used could be important in interpreting results of research on carabids using pitfall traps. In this study we compared the effects of six pitfall trap preservatives on collections of midwestern forest ground beetles.

MATERIALS AND METHODSThe study was conducted in a deciduous forest in McDonough Co., Illinois

from May to October 2005. The site was located at 40.4973° N and 90.5993° W. Dominant tree species consisted of white oak (Quercus alba L.), northern red oak (Quercus rubra L.), black oak (Quercus velutina Lam.), shagbark hickory (Carya ovata (Miller) K. Koch.), and red maple (Acer rubrum L.). Plant nomenclature follows that of Gleason and Cronquist (1991). Sixty pitfall traps were deployed in ten rows of six traps each. Each trap consisted of two 473 ml plastic cups (Solo®, Urbana, IL) one nested inside the other so that the inner cup could be removed during collections and replaced with a fresh one with minimal disturbance to the trap site. The diameter of the cup opening was 9.3 cm. Traps were placed so the trap rim was flush with the ground, and efforts were made to return surrounding soil and litter to former conditions. Traps within rows were five meters apart and rows were six meters apart. In each row, each trap was filled with approximately 150 ml of one of six preservatives: 1) 5% acetic acid solution, 2) distilled water, 3) 70% ethanol, 4) 50% ethylene glycol solution (using Prestone® antifreeze), 5) 50% propylene glycol solution (using Sierra® antifreeze), and 6) 10% saline solution (10 g rock salt per 90 ml water). Distilled water was used as the solvent/diluent for all solutions. Traps were operated for eight 5-day trapping periods: 22 – 27 May, 12 – 17 June, 1 – 6 July, 22 – 27 July, 8 – 13 August, 23 – 28 August, 10 – 15 September, and 30 September – 5 October. For each trapping period, fresh preservative was used, and positions of the six treatments were randomly assigned within each row, with the caveat that each treatment was assigned to each posi-tion at least once and not more than twice over the course of the study. This was done to control for possible trap location effects. A drop of unscented detergent was placed in each trap to reduce surface tension. Traps were collected at the end of each trapping period, and carabids were collected, pinned, and identified using a synoptic reference collection of local ground beetle species. Instances of trap disturbance (trap pulled out of the ground and/or mutilated) were noted.

Species richness and Simpson’s diversity indices were calculated for each preservative. Species richness is associated with sample size, so rarefaction was used in comparing species richness of different preservatives. Rarefaction provides an estimate of the expected number of species for a given sample size (Krebs 1999). The University of Alberta Department of Biology online rarefac-tion calculator (U of A 2007) was used in these analyses. Differences in num-bers of beetles collected among treatments were analyzed using permutational multivariate analysis of variance (PERMANOVA – Anderson 2001, McArdle and Anderson 2001). A one-way design was used, with beetle numbers summed across dates, and rows serving as replicates. Beetle numbers were expressed as numbers per trap to compensate for instances of trap disturbance. Analyses were done for all species collectively and for each of eleven species that produced the greatest number of captures. Results of paired comparisons among treat-ments were evaluated using the Bonferroni method to control for Type I error associated with multiple tests (Sokal and Rohlf 1995). This produced a threshold P-value of 0.0033 (0.05 divided by 15 – the number of paired comparisons among 6 treatments) for evaluating significance of paired comparisons.


RESULTSA total of 819 carabids, representing 33 species and 19 genera, were

collected over the course of the study (Table 1), resulting in a capture rate of 0.35 beetles/trap/day. Mean number collected per trapping period (± SE) was 102.38 ± 22.59, with a low of 29 for the 13 August collection and a high of 217 for the 6 July collection. Thirteen instances of trap disturbance occurred over the course of the study, for a disturbance rate of 2.7%. There was no apparent association between trap disturbance and treatment. Each treatment had at least one disturbance, and none more than three disturbances. Species richness per treatment ranged from a low of 17 for saline to a high of 23 for ethylene glycol (Table 1). Rarefaction estimates (Table 1) indicated that observed spe-cies richness values did not differ from expected for any of the preservatives, based on 95% confidence intervals. Simpson’s diversity indices ranged from a low of 0.86 for acetic acid and distilled water to a high of 0.91 for ethylene glycol (Table 1). Mean number of carabids collected per treatment (± SE) was 136.50 ± 14.39, with a low of 83 total carabids collected in saline and a high of 175 in acetic acid (Table 1). Numbers of carabids collected differed significantly among treatments (Table 2; F = 5.632; df = 5, 54; P = 0.0002). Saline produced significantly fewer captures than did all other preservatives except distilled water, and distilled water produced significantly fewer captures than did acetic acid (Fig. 1; P < 0.0033, each comparison).

Eleven species of carabids each produced at least 22 captures (Table 1), and these species comprised 90.7% of total captures. Significant associations between numbers of captures and treatment were seen in four species: Am-phasia interstitialis (Say) (F = 5.173; df = 5, 54; P = 0.0007), Calathus opaculus LeConte (F = 3.754; df = 5, 54; P = 0.0044), Chlaenius nemoralis Say (F = 3.591; df = 5, 54; P = 0.005), and Cyclotrachelus sodalis (LeConte) (F = 2.217; df = 5, 54; P = 0.0494) (Table 2; Fig. 2). Amphasia interstitialis was significantly more abundant in acetic acid and saline than in ethanol, which collected none of this species (P < 0.0033, each comparison). Calathus opaculus was significantly more abundant in ethylene glycol than in either distilled water or saline (P < 0.0033, each comparison). Neither C. nemoralis nor C. sodalis produced significant pairwise comparisons.

DISCUSSIONTraps containing acetic acid and ethylene glycol collected the greatest

numbers of carabids; however, traps containing acetic acid collected relatively low diversity, based on Simpson’s index, whereas those containing ethylene gly-col collected the greatest diversity (Table 1). These results suggest that either preservative would be effective in maximizing numbers of carabids collected, but ethylene glycol may be preferable if high diversity of beetle captures is the goal. Species richness of carabids collected varied among the different preservatives used, but rarefaction results (Table 1) suggested that differences in richness among preservatives can be explained by differences in numbers of individuals collected. Substantial differences among the preservatives were found in total numbers of carabids collected and in numbers of some individual species. Pitfall traps containing acetic acid and ethylene glycol collected 111% and 99% more carabids than did traps containing saline. These results are consistent with those of Scheller (1984) and Holopainen (1992). Scheller (1984) collected 39% more cara-bids with a 5% acetic acid/2% formaldehyde solution than with water in a study in North Zealand, Denmark. It is difficult to ascertain the relative importance of acetic acid vs. formaldehyde on trapping efficiency in Scheller’s (1984) study, but the 5% acetic acid/2% formaldehyde solution collected significantly more carabids than did a 0.5% formaldehyde solution, suggesting that acetic acid may


ble

1. N

umbe

rs, s

peci

es ri

chne

ss, r

aref

actio

n es

timat

es o

f spe

cies

rich

ness

(± S

D),

and

spec

ies

dive

rsity

of c

arab

id b

eetle

s ca

ptur

ed in

pi

tfall

trap

s co

ntai

ning

the

follo

win

g pr

eser

vativ

es: 1

) 5%

ace

tic a

cid

solu

tion

(AA)

, 2) d

istil

led

wat

er (D

W),

3) 7

0% e

than

ol (E

A), 4

) 50%

et

hyle

ne g

lyco

l sol

utio

n (E

G),

5) 5

0% p

ropy

lene

gly

col s

olut

ion

(PG

), an

d 6)

10%

sal

ine

solu

tion

(SA)

. Tr

appi

ng w

as d

one

for e

ight

5-d

ay

trap

ping

per

iods

from

22

May

to 5

Oct

ober

200

5 in

McD

onou

gh C

o., I

llino

is.

Spec

ies

AA

D

W

EA

E

G

PG

SA

To

tal

Pter

ostic

hus

styg

icus

(Say

) 48

35

41

32

33

18

20

7Pt

eros

tichu

s pe

rmun

dus

(Say

) 31

9

24

14

19

9 10

6Pl

atyn

us d

ecen

tis (S

ay)

12

13

8 16

12

16

77

Cyc

lotr

ache

lus

soda

lis (L

eCon

te)

20

5 17

15

7

9 73

Cal

athu

s op

acul

us L

eCon

te

8 5

9 21

12

3

58C

hlae

nius

nem

oral

is S

ay

3 4

17

11

19

2 56

Cyc

lotr

ache

lus

sexi

mpr

essu

s LeC

onte

13

7

6 12

9

1 48

Synu

chus

impu

ncta

tus (

Say)

10

5

5 16

6

4 46

Amph

asia

inte

rstit

ialis

(Say

) 11

4

0 2

3 8

28An

isod

acty

lus

agri

cola

(Say

) 1

5 5

2 7

2 22

Poec

ilus

lucu

blan

dus (

Say)

6

3 2

5 3

3 22

Patr

obus

long

icor

nis (

Say)

3

2 1

2 1

2 11

Chl

aeni

us e

mar

gina

tus S

ay

1 0

4 0

1 1

7C

hlae

nius

tric

olor

Dej

ean

1 1

3 0

2 0

7H

arpa

lus

herb

ivag

us S

ay

1 0

0 2

2 1

6N

otio

philu

s no

vem

stri

atus

(LeC

onte

) 1

2 1

2 0

0 6

Gal

erita

janu

s (F

abri

cius

) 0

0 3

1 1

0 5

Chl

aeni

us p

laty

deru

s Ch

audo

ir

1 1

0 2

0 0

4D

icae

lus

furv

us D

ejea

n 1

1 0

1 0

1 4

Har

palu

s pe

nsyl

vani

cus

(DeG

eer)

0

1 0

3 0

0 4

Cym

indi

s am

eric

anus

Dej

ean

1 0

0 0

0 2

3Sc

aphi

notu

s el

evat

us (H

alde

man

) 0

0 1

0 2

0 3

Chl

aeni

us p

usill

us S

ay

0 1

1 0

0 0

2H

arpa

lus

prot

ract

us C

asey

1

0 1

0 0

0 2

Pent

agon

ica

pict

icor

nis

Bate

s 1

1 0

0 0

0 2

Poec

ilus

chal

cite

s (S

ay)

0 0

0 2

0 0

2Pt

eros

tichu

s pr

aete

rmis

sus

Chau

doir

0

0 0

1 0

1 2


ble

1. C

ontin

ued.

Spec

ies

AA

D

W

EA

E

G

PG

SA

To

tal

Chl

aeni

us p

enns

ylva

nicu

s Sa

y 0

1 0

0 0

0 1

Cic

inde

la s

exgu

ttata

Fab

rici

us

0 1

0 0

0 0

1D

icae

lus

elon

gatu

s Bo

nelli

0

0 0

1 0

0 1

Dic

aelu

s pu

rpur

atus

Bon

elli

0 0

0 1

0 0

1Pt

eros

tichu

s fe

mor

alis

(Kir

by)

0 0

0 1

0 0

1Tr

icho

tichn

us fu

lgen

s (C

siki

) 0

0 1

0 0

0 1

Tota

l 17

5 10

7 15

0 16

5 13

9 83

81

9Sp

ecie

s Ri

chne

ss

21

21

19

23

18

17

Rare

fact

ion

Estim

ate

21.9

± 2

.03

18.6

± 1

.99

20.8

± 2

.03

21.4

± 2

.03

20.3

± 2

.03

17.0

± 1

.93

Sim

pson

’s D

iver

sity

0.

86

0.86

0.

87

0.91

0.

88

0.89


ble

2. R

esul

ts o

f PER

MAN

OVA

s of o

vera

ll an

d sp

ecie

s-sp

ecifi

c car

abid

capt

ures

in p

itfal

l tra

ps co

ntai

ning

the

follo

win

g pr

eser

vativ

es: 1

) 5%

ac

etic

aci

d so

lutio

n (A

A), 2

) dis

tille

d w

ater

(DW

), 3)

70%

eth

anol

(EA)

, 4) 5

0% e

thyl

ene

glyc

ol s

olut

ion

(EG

), 5)

50%

pro

pyle

ne g

lyco

l sol

utio

n (P

G),

and

6) 1

0% s

alin

e so

lutio

n (S

A).

For e

ach

anal

ysis

, Pre

serv

ativ

e df

= 5

, Res

idua

l df =

54,

and

Tot

al d

f = 5

9. T

rapp

ing

was

don

e fo

r eig

ht

5-da

y tr

appi

ng p

erio

ds fr

om 2

2 M

ay to

5 O

ctob

er 2

005

in M

cDon

ough

Co.

, Illi

nois

.

Gro

up

Sour

ce

SS

MS

F P

All B

eetle

s Pr

eser

vativ

e 80

61.4

0 16

12.2

8 5.

63

0.00

02

Re

sidu

al

1546

0.02

28

6.30

Tota

l 23

521.

42

Amph

asia

inte

rstit

ialis

(Say

) Pr

eser

vativ

e 46

479.

63

9295

.93

5.17

0.

0007

Resi

dual

97

038.

48

1797

.01

To

tal

1435

18.1

1

An

isod

acty

lus

agri

cola

(Say

) Pr

eser

vativ

e 11

773.

73

2354

.75

1.10

0.

3892

Resi

dual

11

5124

.23

2131

.93

To

tal

1268

97.9

5

C

alat

hus

opac

ulus

LeC

onte

Pr

eser

vativ

e 37

607.

98

7521

.60

3.75

0.

0044

Resi

dual

10

8193

.87

2003

.59

To

tal

1458

01.8

5

C

hlae

nius

nem

oral

is S

ay

Pres

erva

tive

3927

0.31

78

54.0

6 3.

59

0.00

50

Re

sidu

al

1181

01.4

3 21

87.0

6

Tota

l 15

7371

.75

Cyc

lotr

ache

lus

sexi

mpr

essu

s (Le

Cont

e)

Pres

erva

tive

2379

7.06

47

59.4

1 1.

98

0.08

27

Re

sidu

al

1300

22.1

9 24

07.8

2

Tota

l 15

3819

.25

Cyc

lotr

ache

lus

soda

lis (L

eCon

te)

Pres

erva

tive

2599

7.75

51

99.5

5 2.

22

0.04

94

Re

sidu

al

1266

46.6

1 23

45.3

1

Tota

l 15

2644

.36

Plat

ynus

dec

entis

(Say

) Pr

eser

vativ

e 11

615.

12

2323

.02

1.09

0.

3706

Resi

dual

11

5586

.73

2140

.50

To

tal

1272

01.8

5

Po

ecilu

s lu

cubl

andu

s (S

ay)

Pres

erva

tive

4018

.14

803.

63

0.35

0.

9113

Resi

dual

12

3060

.38

2278

.90

To

tal

1270

78.5

2


ble

2. C

ontin

ued.

Gro

up

Sour

ce

SS

MS

F P

Pter

ostic

hus

perm

undu

s (S

ay)

Pres

erva

tive

1856

1.99

37

12.4

0 1.

67

0.12

04

Re

sidu

al

1200

15.4

5 22

22.5

1

Tota

l 13

8577

.44

Pter

ostic

hus

styg

icus

(Say

) Pr

eser

vativ

e 94

98.6

4 18

99.7

3 1.

27

0.23

66

Re

sidu

al

8109

0.85

15

01.6

8

Tota

l 90

589.

49

Synu

chus

impu

ncta

tus (

Say)

Pr

eser

vativ

e 77

44.5

4 15

48.9

1 0.

59

0.74

53

Re

sidu

al

1421

29.5

0 26

32.0

3

Tota

l 14

9874

.04


have contributed to the increased number of carabids collected. In a study of carabids in central Finland, Holopainen (1992) collected 58% more carabids in pitfall traps containing ethylene glycol than in traps containing water; however, Lemieux and Lindgren (1999) found no difference between ethylene glycol and brine trapping efficiency of carabids in British Columbia, Canada.

Because it is relatively inexpensive, has good preservative properties, and is widely available, ethylene glycol is currently a commonly used pitfall trap preservative (Woodcock 2005), however, concerns about its attractiveness and toxicity to mammals, including pets (Beasley 1985, Marshall and Doty 1990), have led some workers to consider the less toxic propylene glycol as an alternative (Hall 1991). In our study, traps containing ethylene glycol accounted for only two of 13 total trap disturbances, suggesting that this preservative, compared with the others, was not unusually attractive to mammals.

Significant differences in captures among preservatives were found for four ground beetle species, and two (A. interstitialis and C. opaculus) produced significant pairwise comparisons (Fig. 2). Differences among preservatives in numbers and species of carabids captured may result from differential attraction/repellency, differences in escape rates, or some combination of these factors. In our study, pitfall traps containing the four preservatives that appeared to pro-duce the strongest volatiles (acetic acid, ethanol, ethylene glycol, and propylene glycol) collected much greater numbers of carabids than did those containing distilled water or saline. This suggests that chemical attraction could have played a role in these differences, since many insects are known to rely heav-ily on semiochemical perception; however, the former four preservatives also

Figure 1. Mean numbers of carabids collected per trap per trap row (± SE) in pitfall traps containing the following preservatives: 1) 5% acetic acid solution, 2) distilled wa-ter, 3) 70% ethanol, 4) 50% ethylene glycol solution, 5) 50% propylene glycol solution, and 6) 10% saline solution. Trapping was done for eight 5-day trapping periods from 22 May to 5 October 2005 in McDonough Co., Illinois. Means with the same letter were not significantly different at the 0.0033 level derived using the Bonferroni method.


may kill the beetles more quickly, reducing the probability of escape. Saline in particular may also provide enough buoyancy to allow a greater possibility of escape, especially if large-sized or large numbers of beetles create a surface layer on which subsequently captured beetles can crawl. The tendency of arthropods to float in brine may contribute to lower numbers of ground beetle genera and spider individuals captured in that preservative (Schmidt et al. 2006), however, relatively high numbers of A. interstitialis were collected in saline in our study (Fig. 2), suggesting that this species may be attracted to saline or may not be as capable of escape as are other species. Trap material may also have an effect. Glass provides fewer abrasions and less traction for carabids to escape than does plastic. This is probably not a problem with fast-killing preservatives (Luff 1975). Waage (1985) found no difference in trapping efficiency between plastic and glass pitfall traps containing preservative, but empty glass traps had higher catches than empty plastic ones. Saline may not be a desirable preservative to use with plastic pitfall traps if beetles have long survival times and float on the surface, thus having greater opportunities to escape.

It is also possible that differences in attraction or repellency may result from secondary volatiles produced by non-target organisms, such as other ar-thropods, gastropods, or earthworms collected in the traps. This would probably become more of a factor in studies employing longer trapping intervals than our relatively short 5-day trapping periods, particularly for distilled water and saline, which would allow more rapid decomposition to take place. Dilution of

Figure 2. Mean numbers of four species of carabids collected per trap per trap row (± SE) in pitfall traps containing the following preservatives: 1) 5% acetic acid solution, 2) distilled water, 3) 70% ethanol, 4) 50% ethylene glycol solution, 5) 50% propylene glycol solution, and 6) 10% saline solution. Trapping was done for eight 5-day trapping periods from 22 May to 5 October 2005 in McDonough Co., Illinois. For A. interstitialis and C. opaculus, means with the same letter were not significantly different at the 0.0033 level derived using the Bonferroni method. For C. nemoralis and C. sodalis, no pairwise comparisons were significant.


preservative by rainwater would also contribute to rapid decomposition. In our study, greater than trace amounts of rain occurred during two trapping periods, the night of 27-28 August and the night of 13-14 September. In both cases, the rain occurred near the end of the trapping period, and trap liquid levels increased by approximately 1 cm in each case. Under conditions in which preservative dilution was greater or occurred earlier in the trapping period, significant decomposition probably would have taken place; however, decompos-ing carrion may have little or no effect on ground beetle captures. Greenslade (1964) found that baiting pitfall traps with meat or carrion did not influence collections of carabids.

Carabids collected during this study were generally in good condition, although, compared with the other preservatives, some noticeable softening of specimens collected in distilled water and saline did occur. This indicates that, under environmental conditions similar to those in which this study was done, these two preservatives would probably not be suitable for studies incorporating long trapping periods between collections. Because ethanol tends to evaporate and become diluted relatively rapidly, it would probably be undesirable in such studies as well. Acetic acid is also known to soften ground beetle specimens after several weeks, and would probably produce poor specimens if left for extended periods (anonymous reviewer, pers. comm.), however, beetles collected in these latter two preservatives in our study ap-peared to be in good condition, probably due to the short trapping periods and primarily shaded conditions.

Type of preservative used is also an important consideration in studies requiring isolation of DNA from trapped arthropods. Gurdebeke and Maelfait (2002) found that 70% ethanol was superior to 4% formaldehyde and a 1:1 mix-ture of acetic acid:TE buffer (Tris + EDTA) in short term laboratory storage of the amaurobiid spider Coelotes terrestris (Wider) for DNA analysis, however, for field collection using pitfall traps collected weekly, they found that 96% ethanol was superior to either 85% or 75% ethanol. In a study of the population genet-ics of the ground beetle Chlaenius platyderus Chaudoir, relatively undegraded DNA has been successfully extracted from beetles captured in pitfall traps containing undiluted propylene glycol antifreeze (M. A. Romano, pers. comm.). Beetles in that study were collected twice weekly and transferred to 95% ethanol after collection. Based on these results, at least two of the preservatives used in this study (ethanol and propylene glycol), if used undiluted or nearly so, are potentially useful in ground beetle population genetics studies requiring DNA extraction and analysis.

Results of this study show that the type of preservative used in pitfall trapping studies can affect collections of carabids and that these effects can be species-specific. This indicates that estimates of ground beetle abundance, species composition, and diversity may not be comparable among studies using different pitfall trap preservatives. Ground beetle researchers should carefully consider the goals of their studies when choosing a pitfall trap preservative.

ACKNOWLEDGMENTSWe thank Rachel Knight for help with installing pitfall traps and with

collecting and processing samples. Special thanks also to anonymous reviewers for valuable comments on the manuscript, insights regarding statistical analyses, information on PERMANOVA, and helpful references.

LITERATURE CITEDAnderson, M. J. 2001. A new method for non-parametric multivariate analysis of variance.

Austral Ecol. 26: 32-46.


Beasley, V. R. 1985. Diagnosis of ethylene glycol (antifreeze) poisoning. Feline Practice 15: 41-46.

Curtis, D. J. 1980. Pitfalls in spider community studies (Arachnida, Araneae). J. Arach-nol. 8: 271-280.

Digweed, S. C., C. R. Currie, H. A. Carcamo, and J. R. Spence. 1995. Digging out the “digging-in effect” of pitfall traps: influences of depletion and disturbance on catches of ground beetles. Pedobiologia 39: 561-576.

Gleason, H. A., and A. Cronquist. 1991. Manual of Vascular Plants of Northeastern United States and Adjacent Canada, 2nd ed. New York Botanical Garden, New York.

Greenslade, P. J. M. 1964. Pitfall trapping as a method for studying populations of Carabidae (Coleoptera). J. Anim. Ecol. 33: 301-310.

Greenslade, P. J. M. 1973. Sampling ants with pitfall traps: digging in effects. Insectes Soc. 20: 343-353.

Gurdebeke, S., and J.-P. Maelfait. 2002. Pitfall trapping in population genetics studies: finding the right “solution.” J. Arachnol. 30: 255-261.

Hall, D. W. 1991. The environmental hazard of ethylene glycol in insect pit-fall traps. Coleopt. Bull. 45: 193-194.

Halsall, N. B., and S. D. Wratten. 1988. The efficiency of pitfall trapping for polyphagous predatory Carabidae. Ecol. Entomol. 13: 293-299.

Hammond, P. M. 1990. Insect abundance and diversity in the Dumoga-Bone national park, N. Sulawesi, with special reference to the beetle fauna of lowland rain forest in the Toraut region, pp. 197-254. In W. J. Knight and J. D. Holloway (eds.), Insects and the Rainforests of South East Asia (Wallacea). The Royal Entomological Society of London, London.

Holopainen, J. K. 1990. Influence of ethylene glycol on the numbers of carabids and other soil arthropods caught in pitfall traps, pp 339-341. In N. E. Stork (ed.), The Role of Ground Beetles in Environmental and Ecological Studies. Intercept, Hamp-shire, UK.

Holopainen, J. K. 1992. Catch and sex ratio of Carabidae (Coleoptera) in pitfall traps filled with ethylene glycol or water. Pedobiologia 36: 257-261.

Holopainen, J. K., and A.-L. Varis. 1986. Effects of a mechanical barrier and formalin preservative on pitfall catches of carabid beetles (Coleoptera, Carabidae) in arable fields. J. Appl. Entomol. 102: 440-445.

Honêk, A. 1988. The effects of crop density and microclimate on pitfall trap catches of Carabidae, Staphylinidae (Coleoptera), and Lycosidae (Araneae) in cereal fields. Pedobiologia 32: 233-242.

Ings, T. C., and S. E. Hartley. 1999. The effect of habitat structure on carabid commu-nities during the regeneration of a native Scottish forest. For. Ecol. Manage. 119: 123-136.

Krebs, C. J. 1999. Ecological Methodology, 2nd ed. Benjamin/Cummings, Menlo Park, CA.

Lemieux, J. P., and B. S. Lindgren. 1999. A pitfall trap for large-scale trapping of Cara-bidae: comparison against conventional design, using two different preservatives. Pedobiologia 43: 245-253.

Lövei, G. L., and K. D. Sunderland. 1996. Ecology and behavior of ground beetles (Co-leoptera: Carabidae). Annu. Rev. Entomol. 41: 231-256.

Luff, M. L. 1968. Some effects of formalin on the numbers of Coleoptera caught in pitfall traps. Entomol. Mon. Mag. 104: 115-116.

Luff, M. L. 1975. Some features influencing the efficiency of pitfall traps. Oecologia 19: 345-357.


Marshall, D. A., and R. L. Doty. 1990. Taste responses of dogs to ethylene glycol, pro-pylene glycol, and ethylene glycol-based antifreeze. J. Am. Vet. Med. Assoc. 12: 1599-1602.

McArdle, B. H., and M. J. Anderson. 2001. Fitting multivariate models to community data: a comment on distance-based redundancy analysis. Ecology 82: 290-297.

McCravy, K. W., and J. E. Willand. 2005. Islands in the beans: diversity of ground beetles (Coleoptera: Carabidae) inhabiting west-central Illinois coal mine remnants. Am. Entomol. 51: 234-236.

Niemelä, J., J. R. Spence, and D. H. Spence. 1992. Habitat associations and seasonal activity of ground-beetles (Coleoptera, Carabidae) in Central Alberta. Can. Entomol. 124: 521-540.

Scheller, H. V. 1984. Pitfall trapping as the basis for studying ground beetle (Carabidae) predation in spring barley. Tidsskr. Planteavl. 88: 317-324.

Schmidt, M. H., Y. Clough, W. Schulz, A. Westphalen, and T. Tscharntke. 2006. Capture efficiency and preservation attributes of different fluids in pitfall traps. J. Arachnol. 34: 159-162.

Sokal, R. R., and F. J. Rohlf. 1995. Biometry, 3rd ed. Freeman and Co., New York.Thiele, H-U. 1977. Carabid Beetles in Their Environments. Springer-Verlag, Berlin/

Heidelberg.(U of A) University of Alberta. 2007. Rarefaction calculator. Department of Biology,

University of Alberta. Available at: <http://www.biology.ualberta.ca/jbrzusto/rar-efact.php#Calculator>

Villa-Castillo, J., and M. R. Wagner. 2002. Ground beetle (Coleoptera: Carabidae) species assemblage as an indicator of forest conditions in northern Arizona ponderosa pine forests. Environ. Entomol. 31: 242-252.

Waage, B. E. 1985. Trapping efficiency of carabid beetles in glass and plastic pitfall traps containing different solutions. Fauna Norv., Ser. B 32: 33-36.

Willand, J. E., and K. W. McCravy. 2006. Variation in diel activity of ground beetles (Coleoptera: Carabidae) associated with a soybean field and coal mine remnant. Gt. Lakes Entomol. 39: 141-148.

Woodcock, B. A. 2005. Pitfall trapping in ecological studies, pp. 37-57. In S. R. Leather (ed.), Insect Sampling in Forest Ecosystems. Blackwell Publishing, Malden, MA.

Work, T. T., C. M. Buddle, L. M. Korinus, and J. R. Spence. 2002. Pitfall trap size and capture of three taxa of litter-dwelling arthropods: implications for biodiversity stud-ies. Environ. Entomol. 31: 438-448.


PARASITISM OF UROPHORA AFFINIS (DIPTERA: TEPHRITIDAE) BY APROSTOCETUS SP. (HYMENOPTERA: EULOPHIDAE)

IN MICHIGANJordan M. Marshall1

Urophora affinis Frfld. and U. quadrifasciata (Meig.) (Diptera: Tephriti-dae) are Eurasian gallicolous fruit flies introduced to North America in 1972 as biological control agents for Centaurea biebersteinii DC (spotted knapweed, Asteraceae, = C. maculosa auct. non Lam.) (Harris 1980). Through natural dispersal and numerous introductions, both Urophora species have become distributed throughout the introduced range of C. biebersteinii (Foote et al. 1993, Lang et al. 1997, Mays and Kok 2003). Use as biological control agents for C. biebersteinii focused on the diversion of energy from seed production to the development of Urophora spp. larvae within galls (Harris 1980). In the introduced range of U. affinis and U. quadrifasciata, mortality of these species has occurred as a result of predation by bird and mammal species, as well as parasitism by Pteromalus cardui (Erdös) (Hymenoptera: Pteromalidae) (Story et al. 1995, Pearson 1999, Marshall et al. 2004).

Aprostocetus sp. (Hymenoptera: Eulophidae) was initially observed emerg-ing from C. biebersteinii seed heads in rearing during Feburary 2008. Typically the subfamily Tetrastichinae, containing the genus Aprostocetus Westwood, is endoparasitic of eggs, larvae, or pupae of Coleoptera, Diptera, Hymenoptera, and Lepidotera, with a distinct association with gall inducing hosts (Noyes 2003, Yegorenkova et al. 2007). Species within Aprostocetus, approximately 670, are distributed globally (Graham 1987, LaSalle 1994, Yegorenkova et al. 2007).

Centaurea biebersteinii seed heads were collected from three sites in Livingston County, MI, on 18 February 2008. Sixty seed heads from each site were randomly selected and placed into 8 dram plastic shell vials for rearing. Vials were a third filled with wet sand, topped with a layer of dry sand, and were capped with cotton fabric. Each vial contained 2 C. biebersteinii seed heads (30 vials/site), stored at 80° C at 45 percent humidity, and were checked every 2 days until emergence began. Following initial emergence, vials were checked daily and Aprostocetus sp. adults were removed and sexed. Adult Urophora species were identified using Foote et al. (1993). Chi-square tests were used to test for the independence of U. affinis from the occurrence of Aprostocetus sp. A t-test was used to test for differences in U. affinis emergence in the presence of Aprostocetus sp.

Aprostocetus sp. first emerged after 14 days. Seed heads were left in vials for an additional 10 weeks, which resulted in 34.4 percent of vials producing a total of 373 adult Aprostocetus sp. All emergence of Aprostocetus sp. occurred during days 14-17. In vials that produced adult Aprostocetus sp., 12.03 ± 9.04 individuals emerged per vial with a sex ratio of 0.36 males/females. A total of 36 adult U. affinis emerged from 26.7 percent of vials with a sex ratio of 0.89 males/females. In vials that produced adult U. affinis, 1.50 ± 0.83 individuals emerged per vial. The presence of U. affinis was independent of the presence of Aprosto-cetus sp. individuals (χ2 = 0.404, df = 1). There was no significant difference in the count of U. affinis in vials with and without Aprostocetus sp. emergence (t = 0.672, df = 88). In addition to Aprostocetus sp. and U. affinis, three P. cardui females and 10 U. quadrifasciata (6 males and 4 females) emerged.

Seed heads not placed into rearing were dissected and inspected for U. af-finis and U. quadrifasciata galls. Galls produced by these two Urophora species are structurally different, with U. affinis producing woody, lignified galls and

1Michigan Technological University, Cooperative Emerald Ash Borer Project, 5936 Ford Ct. Suite 200, Brighton, MI, 48116. (e-mail: [email protected]).


U. quadrifasciata producing papery, non-lignified galls (White and Korneyev 1989, Burkhardt and Zwölfer 2002). Only galls produced by U. affinis were en-countered. A sample of 15 U. affinis galls was dissected and 8 contained multiple Aprostocetus sp., with a mean parasitism rate of 8.36 ± 4.07 Aprostocetus sp. individuals per U. affinis gall.

The gregarious endoparasitism of Aprostocetus sp. on U. affinis caused mortality in U. affinis at the sites in Livingston County, MI, where seed heads were collected. While this mortality did reduce the mean number of U. affinis individuals emerging within vials, it did not create significant differences in the vials with and without Aprostocetus sp. and did not significantly influence the presence of U. affinis individuals. With this first record of Aprostocetus sp. parasitizing U. affinis, further investigations into host selection and the geo-graphic distribution of this parasitoid within the introduced ranges of Urophora spp. are warranted.

ACKNOWLEDGMENTSThe author would like to thank Roger A. Burks (University of California,

Riverside) for the identification of Aprostocetus sp. and Jonathan P. Lelito (The Pennsylvania State University) for lab assistance. Voucher specimens are held in the personal collection of the author.

LITERATURE CITEDBurkhardt, B., and H. Zwölfer. 2002. Macro-evolutionary tradeoffs in the tephritid Uro-

phora: benefits and costs of an improved plant gall. Evol. Ecol. Research 4: 61-77.Foote, R. H., F. L. Blanc, and A. L. Norrbom. 1993. Handbook of the Fruit Flies (Dip-

tera: Tephritidae) of America North of Mexico. Cornell University Press, Ithaca, New York.

Harris, P. 1980. Establishment of Urophora affinis Frfld. and U. quadrifasciata (Meig.) (Diptera: Tephritidae) in Canada for the biological control of diffuse and spotted knapweed. Z. Angew. Entomol. 89: 504-514.

Graham, M. W. R. de V. 1987. A reclassifi cation of the European Tetrastichinae (Hy-1987. A reclassification of the European Tetrastichinae (Hy-menoptera: Eulophidae), with a revision of certain genera. Bull. Brit. Mus. (Nat. Hist.), Entomol. Series 55:1-392.

Lang, R. F., R. D. Richard, and R. W. Hansen. 1997. Urophora affinis and U. quadrifasciata (Diptera: Tephritidae) released and monitored by USDA, APHIS, PPQ as biological control agents of spotted and diffuse knapweed. Gt. Lakes Entomol. 30: 105-113.

LaSalle, J. 1994. North American genera of Tetrastichinae (Hymenoptera: Eulophidae). J. Nat. Hist. 28: 109-236.

Marshall, J. M., R. A. Burks, and A. J. Storer. 2004. First host record for Pteromalus cardui (Hymenoptera: Pteromalidae) on Urophora quadrifasciata (Diptera: Tephriti-dae) in spotted knapweed (Centaurea biebersteinii, Asteraceae) in Michigan, U.S.A. Entomol. News 115: 273-278.

Mays, W. T., and L. T. Kok. 2003. Population dynamics and dispersal of two exotic bio-logical control agents of spotted knapweed, Urophora affinis and U. quadrifasciata (Diptera: Tephritidae), in southwestern Virginia from 1986 and 2000. Biological Control 27: 43-52.

Noyes, J. S. 2003. Universal Chalcidoidea Database. The Natural History Museum, London. Available at: <http://www.nhm.ac.uk/entomology/chalcidoids/>

Pearson, D. E. 1999. Deer mouse predation on the biological control agent, Urophora spp., introduced to control spotted knapweed. Northwest. Nat. 80: 26-29.


Story, J. M., K. W. Boggs, W. R. Good, L. J. White, and R. M. Nowierski. 1995. Cause and extent of predation on Urophora spp. larvae (Diptera: Tephritidae) in spotted knapweed capitula. Environ. Entomol. 24: 1467-1472.

White, I. M., and V. A. Korneyev. 1989. A revision of the western Palaearctic species of Urophora Robineau-Desvoidy (Diptera: Tephritidae). Systematic Entomol. 14: 327-374.

Yegorenkova, E. N., Z. A. Yefremova, and V. V. Kostjukov. 2007. Contributions to the knowledge of tetrastichine wasps (Hymenoptera, Eulophidae, Tetrastichinae) of the Middle Volga Region. Entomol. Review 87: 1180-1192.


117 Valleyview Dr., Streator, IL 61364-1114.

ARTHROPODS UTILIZING STICKY INFLORESCENCES OF CIRSIUM DISCOLOR AND PENSTEMON DIGITALIS

Patricia A. Thomas1

ABSTRACTCirsium discolor (Muhl) Spreng (Asteraceae) and Penstemon digitalis Nutt.

(Scrophulariaceae) produce sticky material only in their inflorescences. While there is a wealth of printed information concerning such sticky traps occurring in other parts of plants, there is relatively little about those specifically in inflo-rescences. In order to determine whether sticky traps in the inflorescences of these two plant species defend against seed predators and other herbivores and predators, it was necessary to discover what arthropods use them. Literature search revealed very little about arthropods associated with C. discolor, and nothing about those associated with P. digitalis. Observations showed that, for both plant species, pollinators do not come in contact with the traps, and each plant has several seed predators able to successfully avoid the traps. Several predatory arthropods occur on C. discolor. Two of them, a minute pirate bug and a small salticid spider, seem to glean from its sticky traps. A theridiid spider occasionally builds its web in P. digitalis inflorescences, but was not seen to glean from sticky traps. An undescribed pteromalid parasitizes one of the seed predators of P. digitalis. Ants and aphids are deterred by the traps.

____________________

Hundreds of plant species, in many families, catch insects in sticky traps formed by mucilaginous or resinous secretions. However, in Gray’s Manual of Botany (Fernald 1950) I found only 68 species, divided among 14 families, that have such traps only in their inflorescences. Among these were two species found in Illinois: Foxglove penstemon (Penstemon digitalis Nutt.: Scrophulariaceae) and Field thistle (Cirsium discolor (Muhl) Spreng: Asteraceae).

The function of sticky traps in plants has been debated since Darwin (1875) argued that trapped insects might enhance plant nutrition via direct digestion and absorption. Kerner (1878) countered that their main function was to de-fend against creeping insects. Willson et al. (1983) theorized that when these sticky traps occur in inflorescences they defend against seed predators. The attraction of predators that might remain and defend the plant was suggested by work on carnivorous plants (Lloyd 1942) and extrafloral nectaries (Inouye and Taylor, Jr. 1979). Eisner and Aneshansley (1983) suggested trapped insects might decompose when they were washed to the ground by rain, the products of their decomposition then being absorbed by the plants. From 1980 through 1987 I investigated these four theories: direct nutrition, direct defense against seed predators and other herbivores, indirect defense by attracting predators, and indirect nutrition after decomposition of insects.

The theories of Darwin and of Eisner and Aneshansley require no knowl-edge of the specific insects that come in contact with inflorescences with sticky traps. However, to investigate the defense theories it was necessary to know what insects and other arthropods use the inflorescences, and how they are able to overcome sticky-trap barriers. Behavioral or other adaptations that allow them to do so would be of considerable interest. Although there is very extensive


literature about insects in relation to plant surfaces (Juniper and Southwood, eds. 1986 provide in depth discussions and references), most research papers I found concerned agricultural plants such as tobacco (Van der Plank and An-derssen 1944), tomatoes (Johnson 1956, Stoner et al. 1968, Rick and Tanksley 1981), cotton (Wannamaker 1957, Stephens 1961), and alfalfa (Johnson, Jr. et al.1980). Kerner (1878) mentioned thistles versus ants, and papers by Lamp (1979), Lamp and McCarty (1978), and Rees (1977), considered seed predators of two thistle species other than C. discolor. Only Willson et al. (1983) discussed seed predators of C. discolor. I found no papers about insects utilizing P. digitalis inflorescences. In order to understand insect/plant relationships in the sticky inflorescences of these two plant species, I investigated the arthropods found in their inflorescences. My findings are reported in this paper.

METHODSThis study was done in Trelease Grassland Research Area (TGRA) and

the adjacent Phillips Tract (PT), an area of about 61 hectares, 8 km northeast of Urbana, Illinois. TGRA consists of partially restored prairie that had been farmed until 1949. PT is an old field last farmed in 1950.

Cirsium discolor was found scattered throughout the study area, where it blooms from August into October, each plant bearing many compound inflores-cences. Structures of C. discolor are defended by spines, felting on the under-side of its leaves, and milky sap. Sticky resin is produced by pads on the outer involucral bracts as the small, round buds elongate; these pads are sticky until the inflorescences are dry and brown. The seeds develop rapidly, each with an attached pappus which allows them to be wind-dispersed.

Penstemon digitalis occurred only in TGRA, primarily within 20 to 25 meters of a road running between a wooded area to the north and the open field to the south. Its non-reproductive parts are defended by alkaloids (Lindroth et al. 1986). Each ramet bears only one inflorescence, an indeterminate thyrse. Trichomes occur throughout the inflorescence on peduncles, pedicels, sepals, and the outer surfaces of petals, each trichome secreting and retaining a drop of mucilage on its head. They are present and sticky from the time buds first appear in early spring until all flowers have abscised, by late May or June. The seed capsules are not provided with trichomes. The seeds are small, mature slowly, and are dispersed from the opening tops of the dried capsules in late summer and autumn.

Plants of both species, selected randomly, were observed in the field from the time bud meristems first appeared until seeds were dispersed. The stickiness of the traps and the number of insects they captured were noted, and captured insects were identified at least to order when possible. The insects stuck on P. digitalis trichomes were counted at intervals that allowed old petals to abscise and new flowers to open (2-3 days). Insects stuck on the Cirsium pads were also counted every two or three days; larger insects remained until the inflorescence dried, but very small insects soon sank into the pads and could no longer be identified. I observed and recorded insects visiting, ovipositing, or feeding in the inflorescences and infructescences, collecting some for identification. Ob-servations made during the blooming seasons of the plants (beginning in 1981 and continuing through the 6-year period of concurrent experimental work), amounted to well over 100 hrs. for each plant species.

Inflorescences of both plant species were collected at intervals during their development and flowering, until mature seeds were dispersed. The collected inflorescences were dissected under a stereo-microscope at the University of Il-linois, Champaign-Urbana. Eggs and larvae as well as adult insects present were counted and damage they caused was assessed. Larvae found were reared when possible, for identification. Winter censuses were also done to determine which


insects remained in Penstemon capsules until the following spring. A total of over 630 C. discolor buds, inflorescences and seedheads, and over 7,500 P. digitalis buds, flowers, and seed capsules were dissected, plus 365 P. discolor capsules that had overwintered unopened on 15 stalks from the previous year.

RESULTSResults consist of observations of the insects captured by the sticky traps;

insects using any part of inflorescences for food, including nectar, pollen, struc-tural tissues and seeds, and predatory arthropods including parasitoids.

Insects captured. On P. digitalis I counted up to 918 trapped insects in an inflorescence. Up to 90% of these were small flies, such as midges, blown into the traps by the wind. Fewer, but often larger, insects were trapped on C. discolor bracts (Table 1).

Insects using any part of the inflorescence for food. Cirsium discolor produces copious pollen, which was not fully appreciated in the field because of the numerous insects arriving to eat or collect it. When allowed to open in the lab, however, substantial pollen covered the florets. In the field, I have counted as many as 51 insects of at least eight species on one flower head at one time. Most of these insects landed on the florets but, possibly due to overcrowding on the florets, pollen-eating beetles comprised over 20% of the insects trapped by C. discolor during my study. C. discolor inflorescences also produce nectar and are visited by bees of many species, including: small halictids, honey bees, and small to large bumble bees, as well as many species of butterflies and moths. All of the nectar-users observed landed on the open inflorescence without en-countering the sticky traps.

The principal pollinators of P. digitalis are medium-sized (22 mm) yellow-faced bumble bees (species not identified) that enter the flower directly, fitting precisely into the corolla tube. Within the corolla tube the anthers bend up-ward, so that all brush the upper surface of the hairy thorax of the bees. As the bees enter the next flower, pollen is transferred to the projecting stigma. Larger bumble bees also visit but are unable to enter the flowers. Smaller bees

Table 1. Arthropods trapped by 186 Cirsium discolor inflorescences.

Classification Number trapped Percent of Total

Unidentified 182 27.3Coleoptera 177 26.6Hemiptera 63 9.5Homoptera 42 6.3Ants 52 7.8Other Hymenoptera 57 8.6Lepidopteran larvae* 45 6.8Diptera** 35 5.3Thrips** 6 0.9Arachnida (spiders, mites) 7 1.1Total 666Mean number per inflorescence 3.6

*Lepidopteran caterpillars over-represented; moved out of inflorescences in refrigeratorwhile awaiting further study.**Thrips and very small flies, e.g., probably under-represented; many small, soft-bodied insects sank into the traps and/or became too disintegrated to identify.


of several other species visit the flowers, including: Osmia pumila Cresson and Hoplitis producta Cresson (Megachilidae), Dialictus versatus Robertson and Lasioglossum coriaceum Smith (Halictidae), and honey bees. All land on the inner surface of the petals and do not encounter the sticky trichomes. Occasion-ally one makes a hole near the base of the flower to reach the nectar but is too large to be trapped by the sticky trichomes.

Ants occasionally visit inflorescences of both plant species. They sometimes become trapped on sticky pads but avoid sticky trichomes. When a flower stalk of P. digitalis falls to the ground and flowers can be entered without encounter-ing the trichomes, there is a steady stream of ants collecting nectar.

Aphids rarely colonized C. discolor inflorescences, and readily became trapped when they extended their colonies up the stems onto the inflorescences. No aphids colonized P. digitalis plants or inflorescences during my study. Flying aphids were sometimes caught on the trichomes.

Cirsium discolor was found to have four major seed predators that regu-larly feed in the inflorescences. Destroying over 50% of the potential seeds (Table 2), they were rarely trapped on the sticky pads. They include a tephritid fly and caterpillars of three genera, as follows.

Paracantha culta (Wiedemann) (Diptera: Tephritidae) most often ovipos-ited into the bud meristems. Their early arrival causes very apparent destruc-tion, with deformation of the buds and failure of sticky pads to develop. The larvae (usually one, but up to three, in an inflorescence) eat developing bud meristems, or ovules, seeds, and receptacle, usually completely destroying the seeds. Their occurrence was sporadic. I found none in 78 inflorescences in 1982. In 1983 I found them in two of 60 inflorescences (3.3%). The only year I found them in inflorescences in any numbers was in 1987, when they attacked 19% of the buds. In that year the thistles bloomed early; perhaps P. culta arrived at its usual time, too late to attack meristems before buds had developed, but before the pads had become sticky.

Lobesia carduana (Busck) (Lepidoptera: Tortricidae) caterpillars also arrive early, and make bores or scrapes, covered by a frassy web, in the stem just below the developing inflorescence. They sometimes bore into the base of an inflorescence and eat circumferentially around the receptacle, in which case

Table 2. Percent of Cirsium discolor inflorescences with various seed predators alone or in combination. Combinations not included did not occur. N = 599.

Seed predator Number of Percent of inflorescences inflorescences

H. stypticellum 11 1.8F. tricosa 146 24.4L. carduana 44 7.3P. culta 3 0.5H. stypticellum and F. tricosa 46 7.7H. stypticellum and L. carduana 6 1.0F. tricosa and L. carduana 48 8.0P. culta and H. stypticellum 1 0.2P. culta and F. tricosa 4 0.7P. culta and L. carduana 4 0.7P. culta, H. stypticellum, F. tricosa 2 0.3P. culta, H. stypticellum, L. carduana 1 0.2Totals 316 52.8


the overlying seeds abort. They do not usually eat the seeds. When the frassy web overlies sticky pads, the underlying pads (but not others) appear dry and cleaned-off. The caterpillars may clean off the sticky material or the bores and scrapes may damage the vascular supply to those pads so that sticky material is not produced.

Homeosoma stypticellum Grote (Lepidoptera: Pyralidae) is a common and very destructive seed predator of C. discolor. It usually occurs singly but I have found up to three in an inflorescence. (In the lab, when I put two to-gether, one often ate the other.) H. stypticellum leaves a column of frass and chewed-up pappus as it makes its way from the egg, laid among the florets, to the receptacle. It then eats some developing seeds, but also eats the placental tissue in the receptacle, causing uneaten seeds to abort. Any seeds remaining are often moldy and unable to disperse. The mature caterpillar, too large to be trapped by any remaining sticky material, leaves the inflorescence and probably pupates in the ground.

Two species of Feltia (Lepidoptera: Noctuidae) were found in the inflores-cences but I was able to rear only one species to maturity, Feltia tricosa Lintner. The adult females oviposit among the florets, an unusual place for moths of Noctuinae to oviposit (G. Godfrey, pers. comm.). I have found up to 96 early instars in one flowerhead. As they feed, they seem to produce little damage but a great deal of frass. They spin web among the florets. Their frass is somewhat sticky so that seeds, although not eaten, are often unable to disperse. The small instars usually do not leave the florets and when they do they can become stuck on the bracteal traps. This happened when the inflorescences were kept in the refrigerator for several days before I dissected them and in the field only when the flowerhead was dying unnaturally. In 1987, some disease apparently killed many of the thistle plants and on these dying plants many small Feltia spp. caterpillars left the florets and became trapped. Possibly later instars remain and eat undispersed seeds before leaving the inflorescences to overwinter in the ground. This occurred in one seedhead in the refrigerator but I have not found the caterpillars in old seedheads in the field.

Penstemon digitalis was found to have five major seed predators that regularly feed in the inflorescences destroying over 50% of the potential seeds (Table 3), as follows.

Allophyla atricornis (Meigen) (Diptera: Heleomyzidae) larvae destroyed over 40% of the developing buds. The female oviposits into the buds, leaving a distinctive hole through sepals and petals. Although she oviposits when the trichomes are stickiest, she seldom gets caught by them. For example, 59 in-florescences trapped a total of 8,293 insects, of which only 3 were A. atricornis, while receiving 1,576 A. atricornis ovipositions. When ovipositing the female often flies to a nearby leaf or stem and thoroughly grooms, presumably removing any accumulated mucilage or reapplying some substance to which the trichome

Table 3. Percent of developing buds of P. digitalis lost to herbivores in 1985-1986. N = 6159 buds in 122 inflorescences.

Seed Predator Total number Percent

A. atricornis 2700 43.8Phytomyza sp. 274 4.4P. umbra 214 3.5H. lavana and E. hebesana 124 2.0Total lost to herbivory 3312 53.8


mucilage cannot stick. The egg is deposited into a bud, usually one per bud; if two, the first larva to hatch eats the second. The larva eats the developing pollen from each anther, then the ovules, and finally leaves the bud presumably to pupate in the ground (occasionally one pupates in the corolla tube). Once the style is damaged, the bud fails to open, pollination does not occur, and petals do not dehisce.

An undescribed species, Phytomyza near atripalpis Aldrich (Diptera: Agromyzidae), oviposits in the sepals of large buds or open flowers, tucking the egg under a small flap the female cuts in the abaxial surface of the sepal where there are few sticky trichomes. The larva mines through the sepal into the ovary, where it eats a few developing seeds and then the placenta, causing the rest of the seeds to abort. It pupates in the hole its feeding has produced in the placenta and remains there until eclosing the following spring.

Larvae of three lepidopteran species feed in P. digitalis inflorescences. Hysterosia lavana Busck (Lepidoptera: Cochylidae) and Endothenia hebesana (Walker) (Lepidoptera: Tortricidae) caterpillars eat seeds from inside the seed capsules, which they enter through small holes usually hidden under a sepal. The larvae of both species web their entrance holes shut and remain in the cap-sules until the following spring, when they pupate and eclose. Pyrrhia umbra Hufnagel (Lepidoptera: Noctuidae) caterpillars web flowers together and eat developing capsules from the outside. The Pyrrhia caterpillars I have found were too large to be caught by the sticky traps. In the lab they ate immature capsules but refused P. digitalis leaves and flowers. I have not found the eggs of any of the three lepidopteran species, nor have I found small caterpillars stuck on the traps.

Predatory arthropods feeding in the inflorescences. The situation on C. discolor is very complex, with an array of predators feeding in the different niches the inflorescence provides. Some, including reduviid bugs, mirid bugs, phymatids, phalangids, and thomisid (crab) spiders, are transient opportunists on the florets. Two others, a minute pirate bug and an unidentified small gray jumping spider (Salticidae), appear to live by gleaning from the traps. Ants might glean from the traps, but the only trapped insects I have seen them in-vestigating were living conspecifics. The minute pirate bug, Orius insidiosus (Say) (Anthocoridae) is found occasionally among the florets, where it may eat eggs and small larvae of lepidopterans, but is most often found on the bracts as nymphs and adults. The bug is fairly often trapped on the sticky pads, the salticid spider rarely, if ever. (Perhaps as a spider it knows to avoid stepping in sticky stuff, however, one very small crab spider was found stuck on a trap, straddling an insect also trapped in the sticky material.) The salticid spider’s night-time shelter, a clean web pocket, was often found on top of Lobesia car-duana’s frassy web. In this case the caterpillar was never there. I could not determine whether the borer is prey for the spider or the spider merely uses the caterpillar’s deserted web as a non-sticky base.

Besides the minute pirate bugs, a predatory orange maggot (unidentified dipteran) feeds among the florets, eating cecidomyiid larvae and lepidopteran eggs. Predaceous beetle larvae of at least two families (Cantharidae, Cleridae) occasionally forage among the florets but I have never seen either adults or larvae of these beetles on the outside of the inflorescences.

On P. digitalis the only predatory arthropod I found was an unidentified theridiid spider that occasionally spins its tangleweb in the inflorescences. I have never seen it investigating insects on the traps. Spiders of any species are rarely trapped on the trichomes. Only two spiders were among 14,067 trapped arthropods.

An undescribed pteromalid wasp, Pteromalus sp. (Hymenoptera: Ptero-malidae) parasitizes agromyzid pupae in P. digitalis seed capsules. It may have


either bimodal emergence or more than one generation a year, since when mature capsules were kept in the lab these wasps emerged in November and again in February. In these capsules, only two agromyzids emerged, an unusually low number. Usually when old (previous year) inflorescences were brought into the lab in the spring, pteromalids and agroymyzids emerged in about equal numbers. When pteromalids were first noticed in a plastic bag containing P. digitalis capsules, they were placed in the freezer but remained active 30 minutes later. Perhaps, being so cold tolerant, adults of this pteromalid are able to eclose during the winter and find unparasitized agromyzid pupae in which to oviposit. During my study I did not find this parasitoid visiting P. digitalis flowers.

DISCUSSIONMany species of insects, and a few spiders, were observed using inflo-

rescences of each plant species in spite of the sticky traps. On C. discolor the salticid spider and minute pirate bug appeared to glean from the traps, whereas the others, and all of those on P. digitalis, avoided the sticky materials. Sticky traps may have developed to provide protection to the inflorescences or seeds, but they now seem to protect only against ants and aphids. Apparently coevolu-tion has worked on behalf of the other insects involved, a conclusion enhanced by experiments involving occlusion of sticky traps on these two plant species (Thomas 2003).

Some secretions of trichomes, such as those of Nicotiana sp. (Solanaceae), contain toxins that are rapidly lethal to insects trapped on them (Thurston et al. 1966) but this was not the case here, where insects struggled for prolonged periods of time. Drosophila melanogaster Meigen (Diptera: Drosophilidae) that were caught on P. digitalis in the lab sometimes escaped and though exhibit-ing toxic symptoms, such as lack of coordination, falling onto their backs, etc., were restored to health by careful removal of sticky material from their legs and bodies.

It would be very interesting to investigate the host breadths of the insects that use these two plant species, and whether their behavior varies if they also feed in inflorescences without sticky-traps. Arnett, Jr. (1993) contains general information about feeding habits of the insect families and some of the genera found in this study. Only one species, E. hebesana, the verbena budworm, a horticultural pest, is mentioned with no information about its larval food range. He states there are 6 described species of Paracantha, in the family Tephritidae and mentions larvae of many tephritid species live in developing seeds of composites, some are very destructive pests of fruits, but says nothing specifically about Paracantha species. (Two of the insects associated with P. digitalis, the agromyzid and the pteromalid, are undescribed species.) A great deal of information might be revealed by a literature search, as well as by comparative observations of ovipositing and larval behavior on various larval foods of specific insects.

ACKNOWLEDGMENTSI am very grateful to entomologists at the Illinois Natural History Survey

(NNHS) who identified insects for me: George Godfrey, Wally Laberge, John Bouseman, Steve Heydon, and David Voegtlin; and to Stewart Berlocher, who identified the tephritid flies. Voucher specimens were deposited at the INHS.

LITERATURE CITEDArnett, Ross H., Jr., 1993. American insects: a handbook of the insects of America north

of Mexico. The Sandhill Crane Press, Inc., Gainesville, Florida.


Darwin, C. 1875. Insectivorous plants. John Murray, London.Eisner, T., and D. J. Aneshansley. 1983. Adhesive strength of the insect-trapping glue

of a plant (Befaria racemosa). Ann. Entomol. Soc. Am. 76: 295-298.Fernald, M. L. 1950. Gray’s manual of botany. Eighth edition. American Book, New

York.Inouye, D. W., and O. R. Taylor, Jr. 1979. A temperate region plant-ant-seed predator

system: consequences of extrafloral nectar secretion by Helianthelia quinquenervis. Ecology 60: 1-7.

Johnson, B. 1956. The influence on aphids of the glandular hairs on tomato plants. Plant Pathol. 5: 131-132.

Johnson, K., Jr., E. L. Sorenson, and E. K. Horber. 1980. Resistance of glandular-haired Medicago species to oviposition by alfalfa weevils (Hypera postica). Environ. Ento-mol. 9: 241.

Juniper, B., and R. Southwood (eds.). 1986. Insects and the plant surfaces. Edward Arnold, London.

Kerner, S. 1878. Flowers and their unbidden guests. C. K. Paul, London.Lamp, W. O. 1979. A preliminary study of seed predators of Platte thistle in the Nebraska

sandhills. Trans. Nebr. Acad. Sci. 7:71-74.Lamp, W. O., and M. K. McCarty. 1978. Infestation patterns of the three seed predators

on the Platte thistle in Nebraska. Proc. Nebr. Acad. Sci. Affil. Soc., 88: 15-16.Lindroth, R. L., G. O. Batzli, and S. I. Avildsen. 1986. Lespedeza phenolics and Penste-

mon alkaloids: Effects on digestion efficiencies and growth of voles. J. Chem. Ecol. 12: 713-728.

Lloyd, F. R. 1942. The carnivorous plants. Dover (1976), New York.Rees, N. E. 1977. Impact of Rhinocyllus conicus on thistles in southwestern Montana.

Environ. Entomol. 6: 839-842.Rick, C. M., and S. D. Tanksley. 1981. Genetic variation in Solanus pennelli: comparisons

with two other sympatric tomato species. Pl. Syst. Evol. 139: 11-45.Stephens, S. G. 1961. Further studies on the feeding and oviposition preferences of the

boll weevil (Anthonomus grandis). J. Econ. Entomol. 54: 1085-1090.Stoner, A. K., J. A. Frank, and A. G. Gentile. 1968. The relationship of glandular hairs

on tomatoes to spider mite resistance. Proc. Am. Soc. Hort. Sci. 93: 532-538.Thomas, P. A. 2003. Sticky exudates on the inflorescences of Cirsium discolor (Aster-

aceae) and Penstemon digitalis (Scrophulariaceae) as possible defense against seed predators. Great Lakes Entomol. 36: 112-121.

Thurston, R., W. T. Smith, and B. P. Cooper. 1966. Alkaloid secretion by trichomes of Nicotiana species and resistance to aphids. Ent. Exp. Appl. 9: 428-432.

Van der Plank, J. E., and E. E. Anderssen. 1944. Kromnek disease of tobacco: A promis-ing method of control. Farming in S. Africa 19: 391-394.

Wannamaker, W. K. 1957. The effect of plant hairiness of cotton strains on boll weevil attack. J. Econ. Entomol. 50: 418-423.

Willson, M. F., P. K. Anderson, and P. A. Thomas. 1983. Bracteal exudates in two Cirsium species as possible deterrents to insect consumers of seeds. Am. Midl. Nat. 110: 212-214.


AN ANNOTATED CHECKLIST OF WISCONSIN HANDSOME FUNGUS BEETLES (COLEOPTERA: ENDOMYCHIDAE)

Michele B. Price1 and Daniel K. Young1

ABSTRACTThe first comprehensive survey of Wisconsin Endomychidae was initiated

in 1998. Throughout Wisconsin sampling sites were selected based on habitat type and sampling history. Wisconsin endomychids were hand collected from fungi and under tree bark; successful trapping methods included cantharidin-baited pitfall traps, flight intercept traps, and Lindgren funnel traps. Examina-tion of literature records, museum and private collections, and field research yielded 10 species, three of which are new state records. Two dubious records, Epipocus unicolor Horn and Stenotarsus hispidus (Herbst), could not be con-firmed. Wisconsin distribution, along with relevant collecting techniques and natural history information, are summarized.

____________________

Endomychidae, handsome fungus beetles, is a moderately large family with 1,300 species in about 120 genera worldwide (Strohecker 1986, Lawrence 1991). There are 22 genera and 46 species known to occur in the United States with the majority occurring in the eastern and southeastern regions (Skelley and Leschen 2002). As their common name implies, endomychids can be at-tractively colored and are typically mycophagous, feeding on a wide variety of fungal types from spores and hyphae of microfungi to large Basidiomycetes (Lawrence 1991, Skelley and Leschen 2002). Some species are rather fungal host specific, such as Endomychus biguttatus Say associated with Schizophyllum commune Fries (Leschen and Carlton 1988), Lycoperdina ferruginea LeConte with puffball fungi in the genera Lycoperdon and Calvatia (Pakaluk 1984), and Xenomycetes laversi Hatch with Paxillus atrotomentosus (Batsch ex. Fr.) Fries (Johnson 1986). Holoparamecus caularum (Aubé), Holoparamecus depressus Curtis, Holoparamecus singularis (Beck), Holoparamecus ragusae Reitter, My-cetaea subterranea (Fabricius), and Trochoideus desjardinsi Guerin have been collected in association with stored products, usually feeding on mold (Hinton 1945, Aitken 1975, Bousquet 1990).

Aphorista morosa LeConte has been found in association with a yellow plasmodium of a slime mold (Myxomycetes) (Lawrence 1991). While some endomychid species occur in termite, ant, and bee nests, little is known about their biology (Lawrence 1991, Yanega and Leschen 1994, Skelley and Burgess 1995). Young (1984, 1989) observed four North American species of Endomychi-dae: Aphorista vittata (Fabricius), Danae testacea (Ziegler), L. ferruginea, and Xenomycetes morrisoni Horn, orienting to filter paper baited with cantharidin, a defensive compound produced by most meloid and some oedemerid beetles. The role of cantharidin in endomychid biology is unknown but one conjecture is cantharidin may mimic other terpenoid compounds in nature, such as ter-penoid fungal metabolites that may assist the beetles in locating fungal hosts (Young 1984).

Regardless of the relative abundance and fascinating fungal associations of Endomychidae, natural history and distribution data for species of the western

1Department of Entomology, 445 Russell Labs, 1630 Linden Drive, University of Wis-consin, Madison, Wisconsin, 53706, USA.


Great Lakes region of North America are scarce. Several general beetle reviews and catalogs including endomychids have been conducted for individual states or subdivisions: Dury (1902) for Cincinnati, Ohio; Blatchley (1910) for Indiana; Peck and Thomas(1998) for Florida; or regions: Leng (1920) for North America North of Mexico; Hatch (1961) for the Pacific Northwest; Campbell (1991) for Canada; Downie and Arnett (1996) for the Northeastern United States; Shock-ley and McHugh (2003) for the Great Smoky Mountains National Park. Also, there are a few endomychid specific catalogs of North America: LeConte (1854); Crotch (1873) for the United States; Wickham (1894) for Ontario and Quebec; Strohecker (1986) for America North of Mexico; Arriaga-Varela et al. (2007) for Mexico; and Majka (2007) for the Maritime Provinces of Canada.

In Wisconsin, Rauterberg (1885) recorded a portion of the Coleoptera of Wisconsin collected, “in the vicinity of Milwaukee”, including some endomychid species with brief notes on their abundance (common, rare, or very rare) and how specimens were collected. Wickham (1895) provided a list of Coleoptera from the southern shore of Lake Superior; however, the list did not include en-domychid species from Wisconsin. Until the present study, no comprehensive studies of Wisconsin Endomychidae diversity, life histories, and distributions have been conducted.

MATERIALS AND METHODSThis survey was initiated in 1998 as part of an undergraduate independent

study by the senior author and was continued along with a graduate study of Nitidulidae and Kateretidae of Wisconsin (Price and Young 2006). Colleagues conducting separate insect faunal surveys have and continue to contribute endomychid data. Our paper summarizes data through 2007.

At the onset of this survey, literature records, and museum and private collections were examined to determine which endomychid species had previously been collected in Wisconsin. The following institutional collections were reviewed for Wisconsin records: Field Museum of Natural History (FMNH), Milwaukee Public Museum (MPMC), and University of Wisconsin-Madison Insect Research Collection (WIRC). Several private collections were also examined.

Field survey work focused on historically under-sampled regions and unique Wisconsin habitats, e.g., oak savanna and hemlock forest. A variety of trapping methods were used to collect endomychids, including cantharidin-baited jar traps (Fig. 1) partially buried into the soil, blacklight traps, flight intercept traps, Lindgren funnel traps, and Malaise traps. The following endomychid species were consistantly attracted to cantharidin-baited traps in Wisconsin: A. vittata, D. testacea, and L. ferruginea (Young 1984, 1989). In addition to trap- In addition to trap-In addition to trap-ping, endomychids were hand-collected, mainly from fungi and wood. Specimens were also obtained from sweep net and leaf litter samples.

The following surveys in conjunction with the WIRC also provided Wis-consin specimen records: Wisconsin Department of Natural Resources (WDNR) projects, Hemlock Draw (The Nature Conservancy) and Quincy Bluff (State Natural Area) Surveys, Fort McCoy inventory project, and Necedah National Wildlife Refuge inventory project. Specimen data were entered into the relational biodiversity database software, BIOTATM (Colwell 1996). Voucher specimens have been deposited in the WIRC and additional specimens reside in the personal collections of the contributors to this study.

RESULTSThis survey yielded 10 Wisconsin species in nine genera of Endomychidae

and an additional two dubiously recorded species in two genera. Three species represent new state records not having previously been recorded from the state


in the literature. The arrangement of subfamilies follows the phylogeny proposed by Tomaszewska (2000), with genera and species listed in alphabetical order.

A website has been developed for Wisconsin endomychids. It includes im-ages, distribution maps, and general information for each species. In addition, links are provided to other Wisconsin insect faunal surveys. The website can be accessed at the following address: <http://entomology.wisc.edu/~mbprice/wibeetles/Price/Endo%20Intro.htm>.

For each endomychid species profile, the total number of specimens examined (this number includes specimens with and without ecological data), whether the species is newly reported in Wisconsin, natural history, specific number of species collected in a particular collection event, temporal and Wis-consin geographic distributional records are documented. Species previously recorded in the literature from Wisconsin are followed by the relevant literature reference(s). To simplify county associations, Wisconsin has been divided into nine, 8-county regions (Fig. 2; after Kriska and Young 2002, Hilsenhoff 1995). Life history, phenological, and trapping information pertain solely to adult and larval Wisconsin endomychid records and have been extracted directly from

Figure 1. Disarticulated cantharidin-baited jar trap.


labels accompanying specimens. Thus, in reporting plant and fungal associa-tions, we report the data as indicated by specimen labels only. In some cases the above mentioned are recorded by Latin binomial and author, e.g., under the bark of Populus grandidentata Michaux, while in other cases a common vernacular is used, e.g., on puffball fungus. Many species are active at night when they can be readily collected with a flashlight or headlamp. Those col-lected during the day were typically found in host fungi beneath bark of dead trees or within moist decaying woody material in which fungi were common. Many phenologies are likely artifacts of sampling activity. We still lack sound natural history information for some species in Wisconsin.

LEIESTINAE Thomson, 1863Phymaphora pulchella Newman. NEW STATE RECORD. (81 Wis-

consin specimens examined). Adult specimen collection data included the follow-ing: beneath bark of less than a year old cut log (possibly maple) (5 specimens), under loose bark (8 specimens) under bark of standing dead P. grandidentata abundant with fungus (3 specimens), beneath paper birch bark (2 specimens), under bark of pine (2 specimens), under oak bark (5 specimens), in fungus on

Figure 2. Regional divisions (nine, 8-county areas) of Wisconsin (after Kriska and Young 2002, Hilsenhoff 1995).


fallen tree (1 specimen), and in damp tree-hole litter (3 specimens). We also obtained adult specimens with the following trapping methods: flight intercept traps baited with cantharidin (2 specimens) (bait presumably incidental in this case), Malaise trap (1 specimen), unbaited Lindgren funnel traps (35 speci-mens), as well as, Lindgren funnels baited with cantharidin only (3 specimens), ipsdienol only (4 specimens), both cantharidin and ipsdienol (2 specimens), ethanol (1 specimen), banana and fermenting brown sugar (3 specimens) (all baits presently considered incidental). Aspen and birch dominant forest, mixed conifer-hardwood forest, one-year-old hardwood cut-site, oak and pine forest, beech-dominated northern mesic forest, aspen, oak, Jack pine forest, Pinus resinosa Aiton plantation, oak savanna, southern mesic forest, and northern (wet) mesic forests were observed collection sites. Specimens were active from March to October. Most specimens were collected under bark of fallen trees (25 specimens) and with Lindgren funnel traps (48 specimens): NW: Douglas, Sawyer; NE: Florence, Forest, Marinette, Menominee; WC: Jackson, Monroe; C: Marquette, Wood; SC: Dane, Dodge, Sauk; SE: Racine.

Rhanidea unicolor (Ziegler). (31 Wisconsin specimens examined). Rauterberg (1885) obtained this species with sweep-net and considered it rare. We collected adults beneath bark of a dead log snag near a Lasius Fabricius, ant colony (3 specimens), beneath bark of a log (possibly maple cut less than a year prior) (10 specimens), and in leaf litter (2 specimens). We also obtained adult specimens with the following trapping methods: flight intercept trap (1 specimen), unbaited Lindgren funnel traps (11 specimens), as well as, Lindgren funnel traps baited with banana and fermenting brown sugar (2 specimens) and ipsdienol (2 specimens) (baits considered incidental). Sandy oak bar-rens, southern mesic hardwood forest, oak savanna and plantation with P. resinosa, Pinus strobus Linnaeus, and Picea species were observed collection sites. Specimens were active from April to October. Most specimens were collected under bark of fallen trees (13 specimens) and with Lindgren funnel traps (15 specimens): NE: Shawano; SW: Grant, Richland; SC: Columbia, Dane, Green, Sauk.

ENDOMYCHINAE Leach, 1815Endomychus biguttatus Say. (118 Wisconsin specimens examined).

Rauterberg (1885) obtained this species on young branches of aspen and with sweep-net, and considered the species common. Wisconsin was included in the distribution recorded by Downie and Arnett, Jr. (1996). Adult specimen collec-tion data included the following ecological notations: beneath bark of shagbark hickory snag (1 specimen), beneath bark of P. grandidentata (1 specimen), on pine bark (1 specimen), under bark (1 specimen), under fleshy shelf fungus (1 specimen), crawling on fallen tree (18 specimens), on tree with fungus (2 speci-mens), on small fungus on stick (3 specimens), feeding on fungus on large tooth aspen (4 specimens), on shelf fungus on fallen Populus tree (1 specimen), near sap flow (1 specimen), under leaf litter (1 specimen), in damp tree hole litter at base of tree (1 specimen), and on low vegetation (1 specimen). Ten specimens, including mating pairs, were collected in early September on bark of a fallen tree. One immobile adult specimen was collected at the tip of an Onosmodium molle Michaux inflorescence (1800 h, 80˚F sunny). In April, a larva was collected under bark of a newly cut hardwood. We also obtained adult specimens with the following trapping methods: flight intercept traps (21 specimens), light trap (2 specimens), banana trap (1 specimen), Malaise traps (13 specimens), jug trap (1 specimen), unbaited Lindgren funnel traps (9 specimens), Lindgren funnel traps baited with banana and fermenting brown sugar (3 specimens), and Lindgren funnel traps baited with ipsdienol only (1 specimen) and Lindgren funnel traps baited with cantharidin and ipsdienol (4 specimens) (all baits presently consid-ered incidental). Habitat types included northern (wet and dry) mesic forest,


one-year-old hardwood cut site, dry lime prairie, southern (wet) mesic hardwood forest, oak/pine forest, oak savanna, Pinus banksiana Lambert barrens near lightning struck Quercus spp. snag, and sandy oak barrens. Specimens were active from March to December with the majority collected between July and September. Most specimens were collected on or under bark of fallen trees (32 specimens) and with flight intercept traps (21 specimens) and Lindgren funnel traps (17 specimens): NW: Bayfield, Polk; NC: Vilas; NE: Door, Florence, Forest, Marinette, Shawano; WC: Monroe; C: Marquette, Waupaca, Wood; EC: Fond du Lac; SW: Crawford, Grant, Richland; SC: Dane, Dodge, Green, Lafayette, Rock, Sauk; SE: Jefferson, Ozaukee, Racine, Waukesha.

STENOTARSINAE Gorham, 1873Danae testacea (Ziegler). (>2,000 Wisconsin specimens examined). Ra-

uterberg (1885) collected this species (listed as Mycetina testacea LeConte) under the bark of old logs and considered it rare. Young (1984) obtained this species with cantharidin-baited traps. Ecological data associated with our adult speci-mens included the following: under logs (2 specimens), in leaf litter near fallen tree near fleshy white fungus (6 specimens), in wet moldy leaf litter near fallen tree (3 specimens), leaf litter near fallen tree (8 specimens) and a single specimen was collected at night on leaf of small herbaceous plant. We also obtained adults with the following trapping methods: cantharidin-baited pitfall traps (>1,800 specimens), unbaited (31 specimens) and cantharidin-baited (4 specimens) flight intercept traps, human dung/malt/molasses pitfall traps (6 specimens) (this bait likely incidental), gilled fungus baited pitfall trap (1 specimen), Malaise trap (1 specimen), Lindgren funnel trap baited with ipsdienol (1 specimen), and water-filled pan trap (2 specimens). Collection sites included aspen and birch dominant forest, northern mesic forest (dry), hardwood forest adjacent to lime prairie, southern mesic hardwood forest (dry), oak forest near flood plain, pine and oak forest, sandy oak barrens, and P. resinosa stand. Specimens were ac-tive from March to December. The majority of specimens were collected from pitfall traps baited with cantharidin: NW: Douglas; NE: Florence, Marinette, Shawano; WC: Eau Claire, Jackson, Monroe; C: Adams, Marquette, Waupaca, Waushara, Wood; SW: Grant, Richland, Trempealeau; SC: Dane, Green, Iowa, Lafayette, Sauk; SE: Jefferson, Ozaukee, Racine, Walworth.

LYCOPERDININAE Redtenbacher, 1844Aphorista vittata (Fabricius). (226 Wisconsin specimens examined).

Rauterberg (1885) reported this species as “somewhat common” and Wisconsin was included in the distribution recorded by Strohecker (1986) and Downie and Arnett, Jr. (1996). We collected one adult specimen on a sandy dirt road at the edge of woods and a mating pair was collected in June. We obtained most adults with cantharidin-baited pitfall traps (222 specimens); a single specimen was recovered from a banana trap. Collection sites included northern mesic forest, dry lime prairie, red pine plantation, and oak savanna. Specimens were active from May to August: NW: Douglas; NE: Florence, Marinette; WC: Jackson, Monroe; C: Adams, Wood; SW: Grant; SC: Sauk.

Lycoperdina ferruginea LeConte. (>450 Wisconsin specimens exam-ined). Rauterberg (1885) obtained this species under the bark of old logs and considered it rare. Ackerman and Shenefelt (1973) conducted a general survey of insects associated with macro-fruiting bodies of forest fungi in Wisconsin and recorded L. ferruginea associated with the puffball fungi, Lycoperdon perlatum Persoon and Morganella pyriformis (Schaeffer: Persoon) Krüger and Kreisel (=Lycoperdon pyriforme Schaeffer: Persoon). Young (1984) collected this spe-cies from cantharidin-baited traps in Wisconsin. We collected adult and larval specimens under and in fruiting bodies of puffball fungi (>50 specimens) of the


above-mentioned species and Calvatia species. Larvae were observed from March to April and from September to October in association with puffball fungi. Ecological data associated with our adults include the following: sweep-net (1 specimen), on Geomys bursarius (Shaw) pushup (1 specimen), under log (1 specimen), and in leaf litter near fallen tree (12 specimens). We also obtained adults with the following trapping methods: cantharidin-baited pitfall traps (>350 specimens), cantharidin-baited flight intercept traps (3 specimens), hu-man dung/malt/molasses pitfall trap (1 specimen) (this bait likely incidental), and barrier-pitfall trap near mammal burrow entrance (1 specimen). Collection sites included disturbed grassland, oak savanna, northern mesic forest (dry, old growth), dry sand prairie, edge of lake, old growth hemlock forest, red pine plan-tation, hardwood forest adjacent to a saw mill, sandy oak barrens, and southern mesic hardwood forest. Specimens were active from March to November with the majority of adults from pitfall traps baited with cantharidin: NW: Barron, Burnett; NC: Oneida, Vilas; NE: Florence, Forest, Marinette, Menominee, Shawano; WC: Eau Claire, Jackson, Monroe; C: Adams, Juneau, Marquette, Waupaca, Wood; EC: Manitowoc, Sheboygan; SW: Grant, LaCrosse, Richland; SC: Columbia, Dane, Dodge, Green, Iowa, Lafayette, Sauk; SE: Jefferson, Kenosha, Ozaukee, Walworth, Waukesha.

Mycetina perpulchra (Newman). (107 Wisconsin specimens exam-ined). Rauterberg (1885) collected this species under the bark of old logs and considered it rare. Ecological data from adults we collected included the fol-lowing: on Agaricales fungi (2 specimens), on fungus of downed rotting tree at night (1 specimen), on shelf fungus of decaying tree stump (6 specimens), and in leaf litter (3 specimens). Nine specimens were collected mating on stump covered with fungus in June. We also obtained adults with the following trap-ping methods: unbaited flight intercept traps (52 specimens), flight intercept traps baited with cantharidin (4 specimens), black light (1 specimen), Malaise traps (4 specimens), and a variety of baited Lindgren funnel traps: ipsdienol (4 specimens), banana and fermenting brown sugar (1 specimen), and ethanol (1 specimen); and unbaited Malaise trap (1 specimen). In all cases, including can-tharidin, the bait is presently considered to be incidental and not indicative of a definitive association. Most specimens were collected with the flight intercept traps (56 specimens). Collection sites included aspen and birch dominant forest, northern mesic hardwood forest (dry, wet, old growth, and beech dominated), southern mesic hardwood forest, oak savanna, pine barrens, sandy oak barrens, oak and pine barrens, in ring of declining P. resinosa, and plantation with P. resinosa, P. strobus, and Picea species. Specimens were active from April to October: NW: Bayfield, Douglas, Polk; NC: Ashland, Marathon, Vilas; NE: Florence, Marinette, Menominee; WC: Eau Claire, Jackson, Monroe; C: Juneau, Waupaca, Wood; SW: Grant, LaCrosse, Richland, Trempealeau; SC: Columbia, Dane, Green, Lafayette, Sauk.

MYCETAEINAE Jacquelin du Val, 1857Mycetaea subterranea (Fabricius). Rauterberg (1888) collected this

species with a sweep-net and considered it rare. Pellitteri and Boush (1983) recovered a single specimen in June from a feed mill in southern Wisconsin: SC: Dane.

ANAMORPHINAE Strohecker, 1953Symbiotes duryi Blatchley. NEW STATE RECORD. (9 Wisconsin

specimens examined). We collected adults with unbaited (1 specimen) and cantharidin-baited (1 specimen) flight intercept traps and non-baited Lindgren funnel traps (2 specimens) and Lindgren funnel traps baited with either banana and fermenting brown sugar (3 specimens) or cantharidin (2 specimens). At


this time, all baits are considered to be incidental. Collection sites included dry mesic hardwood forest, northern wet mesic forest, aspen, oak, and jack pine forest, and sandy oak barrens. Specimens were active from May to September: NW: Washburn; NE: Marinette; C: Marquette; SW: LaCrosse, Richland; SC: Sauk; SE: Milwaukee.

Symbiotes gibberosus (Lucas). NEW STATE RECORD. (11 Wis-consin specimens examined). We recovered adults from flight intercept trap (1 specimen) and Lindgren funnel traps (2 specimens) baited with cantharidin and ipsdienol in woods adjacent to Lake Michigan. As with S. duryi, the small number of specimens provides no compelling evidence to suggest a chemical association. In Madison, eight specimens were collected in June with no ad-ditional ecological data. Specimens were active from May to August: SC: Dane; SE: Milwaukee.

The following species represent dubious historical records. No Wisconsin specimens were obtained during the current survey nor were any of Rauter-berg’s specimens bearing these names discovered. Both are almost certainly in error.

STENOTARSINAE Gorham, 1873Stenotarsus hispidus (Herbst). Rauterberg (1885) recorded this spe-

cies from sweep netting and considered it very rare. Stenotarsus hispidus has been reported to occur in southern Indiana and Ohio thus making its presence in Wisconsin slightly plausible; however, this species is more southern in its distribution within North America (Dury 1902, Blatchley 1910).

EPIPOCINAE Gorham, 1873Epipocus unicolor Horn. Rauterberg (1885) obtained this species in au-

tumn in fungus and considered it common. Verified records indicate this species is more southwestern in distribution within North America (Strohecker 1986).

DISCUSSIONIn addition to the two dubious records noted above, the distributional

ranges of several additional endomychid species may extend into Wisconsin. A listing of these follows, along with the state(s) and provinces from which they are currently known in closest proximity to Wisconsin: Bystus ulkei (Crotch) (Indiana), Clemmus minor (Crotch) (Indiana, Illinois), Epipocus punctatus LeConte (Indiana, Illinois), Hadromychus chandleri Bousquet and Leschen (Ontario, Quebec), and Symbiotes impressus Dury (Ohio) (Blatchley 1910, Strohecker 1986, Downie and Arnett, Jr. 1996, Bousquet and Leschen 2002). Except for H. chandleri, these species are more southern in distribu-tion and should any of them extend into Wisconsin they would be expected to occur only in the southern-most portion of the state. H. chandleri ranges from Nova Scotia to southern Ontario and may be restricted to northeastern North America. Natural history information for this species is limited. Two specimens were collected with a flight intercept trap in a red spruce forest and one specimen was collected by shifting a “conifer log” (Bousquet and Leschen 2002). If this endomychid prefers northern wet forests and boreal forest types with species of white and black spruce and other conifers, it could possibly occur in northern Wisconsin.

Within Wisconsin it appears most endomychid species are generally dis-tributed throughout the state in a variety of forested habitats. Several species may show some interesting distribution patterns but sampling bias might also account, at least in part, for the results seen thus far. With 200+ specimens examined, A. vittata may be considered fairly common, yet it is known from only


nine of Wisconsin’s 72 counties. M. perpulchra is fairly widespread through-out the state but lacks records in the east-central and southeastern regions of the state. R. unicolor has been more commonly collected in the southwestern and south-central regions of the state. Although S. duryi is widely distributed throughout the state, it is known from only seven counties. Knowing more about endomychid fungal host specificity and the distributions of preferred fungal hosts would add greatly to such discussions.

The two introduced species, M. subterranea and S. gibberosus, have the fewest county records. Perhaps not surprisingly, both were collected within the vicinity of fairly large cities in the southeastern and south-central parts of Wisconsin.

Although several trapping methods were used to sample endomychids, cantharidin-baited pitfall traps dwarfed all other methods for species known to orient to the compound. More than 1,000 specimens of D. testacea were collected with this method. A. vittata and L. ferruginea were rarely or never collected at flight intercept traps or Lindgren funnel traps, hence pitfall traps baited with cantharidin were a necessary means for collecting these species. While E. biguttatus and M. perpulchra were commonly collected with flight intercept and Lindgren funnel traps, P. pulchella and R. unicolor were commonly collected with mainly Lindgren funnel traps. An awareness of most successful trapping methods for particular species can aid with future studies that monitor habitat management practices and species diversity, especially for forested habitats.

The only previously published list of Wisconsin Endomychidae was over 100 years ago (Rauterberg 1885). The most recent catalog of Endomychidae of America North of Mexico (Strohecker 1986) listed but a single species of En-domychidae from Wisconsin. This updated list of Wisconsin endomychid species along with newly documented associations (e.g., habitat, fungal, floral) offers a new and fairly complete baseline reference point upon which future ecologi-cal monitoring, behavioral, insect-fungus interaction, and other studies might build. This survey complements the growing list of Wisconsin faunal surveys of Coleoptera in progress: Anobiidae, Cantharidae, Cleridae, Lycidae, Meloidae; or completed (Pyrochroidae: Young 1998; Scarabaeoidea: Kriska and Young 2002; Histeridae: Gruber 2003; Mordellidae: Lisberg and Young 2003; Tenebrionidae: Dunford and Young 2004; Silphidae: Katovich, et al. 2005; and Nitidulidae and Kateretidae: Price and Young 2006) in an effort to better understand the “state of the state’s” biodiversity. Such studies provide a long overdue baseline of the Wisconsin beetle fauna and improve our position to evaluate the effects of such major ecosystem perturbations as habitat fragmentation and loss, land use changes, climate change, and impacts of invasive species.

ACKNOWLEDGMENTSSpecial thanks to Theresa Cira for her hard work on the Wisconsin beetle

web pages. We thank our lab colleagues, present and past: Rachel Arango, Devin Biggs, Craig Brabant, Pete DeVries, John Dorshorst, Jim Dunford, Jeff Gruber, Krista Hamilton, Kerry Katovich, Nadine Kriska, Dan Marschalek, Alistair Ramsdale, and Andrew Williams for assistance and contribution of material to this study. Thanks to Phil Pellitteri, University of Wisconsin, Madison for his contribution of literature and to Floyd Shockley at the University of Georgia, Athens for assistance with Symbiotes identifications. We acknowledge cura-tors and staff at the Field Museum of Natural History (Al Newton, Margaret Thayer, Phil Parrillo, and James Boone), Florida State Collection of Arthropods (Mike Thomas and Paul Skelley), Milwaukee Public Museum (Gary Noonan) and University of Wisconsin Madison Insect Research Collection (Steve Krauth) for providing loan material for this study. We thank the Center for Systematic Entomology (Gainesville, Florida), Lois Almon Small Grants Program, and University of Wisconsin-Madison Natural History Museum Council for grant


support. We thank Tom Steele and Karla Ortman at the Kemp Research Station and Betty and Jim Kriska for providing lodging during several field excursions associated with this study. Thomas Meyers, Wisconsin Department of Natural Resources (WDNR) is acknowledged for helping us secure permits to collect within Wisconsin state natural areas. We are grateful to several private land-owners (Richard and Mary Norman, Tom and Eva Wedel) for allowing access to their property to collect and establish trap sites.

LITERATURE CITEDAckerman, J. K., and R. D. Shenefelt. 1973. Organisms, especially insects associated

with wood rotting higher fungi (Basidiomycetes) in Wisconsin forests. Wisc. Acad. Sci., Arts Letters. 61: 185-206.

Aitken, A. D. 1975. Insect travellers I. Coleoptera. Tech. Bull., Ministry of Agriculture, Fisheries and Food. 31:i-xvi + 191pp.

Arriaga-Varela, E., K. W. Tomaszewska, and J. L. Navarrete-Heredia. 2007. A synopsis of the Endomychidae (Coleoptera: Cucujoidea) of Mexico. Zootaxa 1594: 1-38.

Blatchley, W. S. 1910. The Coleoptera or beetles of Indiana: An illustrated descriptive catalogue of the Coleoptera known to occur in Indiana. Bulletin of the Indiana De-partment of Geology and Natural Resources 1: 1-1386.

Bousquet, Y. 1990. Beetles associated with stored products in Canada: an identification guide. Research Branch, Agriculture Canada Publication 1837.

Bousquet, Y. and R. A. B. Leschen. 2002. Description of a New Genus and Species of Endomychidae (Coleoptera: Cucujoidea) from Northeastern North America. Coleopt. Bull. 56(2): 291-298.

Campbell, J. M. 1991. Family Endomychidae (Handsome fungus beetles), pp. 237-239. In Y. Bousquet (ed.), Checklist of beetles of Canada and Alaska, Ottawa Research Branch, Agriculture Canada.

Colwell, R. K. 1996. BIOTA: The biodiversity database manager. Sinauer Associates, Sunderland, MA.

Crotch, G. R. 1873. Synopsis of the Endomychidae of the United States. Trans. Am. Entomol. Soc. 4: 359-363.

Downie, N. M., and R. H. Arnett, Jr. 1996. The beetles of northeastern North America. Volume 2. Sandhill Crane Press, Gainesville, Florida.

Dunford, J. C., and D. K. Young. 2004. An annotated checklist of Wisconsin Darkling Beetles (Coleoptera: Tenebrionidae) with comparisons to the Western Great Lakes fauna. Trans. Am. Entomol. Soc. 130: 57-76.

Dury, C. 1902. A revised list of the Coleoptera observed near Cincinnati, Ohio, with notes on localities, bibliographical references, and description of new species. J. Cincinnati Soc. Nat. Hist. 20: 107-196.

Gruber, J. P. 2003 Hister beetles of Wisconsin. Available at: <http://entomology.wisc.edu/~young/hbhomage/wishis.html>

Hatch, M. H. 1961. The beetles of the Pacific Northwest, Part III: Pselphidae and Diver-sicornia I. University of Washington Publication in Biology 16: 1-503.

Hilsenhoff, W. L. 1995. Aquatic Hydrophilidae and Hydraenidae of Wisconsin (Co-leoptera). I. Introduction, key to genera of adults, and distribution, habitat, life cycle, and identification of species of Helophorus Fabricius, Hydrochus Leach, and Berosus Leach (Hydrophilidae), and Hydraenidae. Great Lakes Entomol. 28: 25-53.

Hinton, H. E. 1945. A monograph of the beetles associated with stored products. Vol. I. British Museum of Natural History, London.


Johnson, P. J. 1986. A description of the late-instar larva of Xenomycetes laversi Hatch (Coleoptera: Endomychidae) with notes on the species’ host and distribution. Proc. Entomol. Soc. Wash. 88: 666-672.

Katovich, K., N. L. Kriska, A. H. Williams, and D. K. Young. 2005. Carrion beetles (Co-leoptera: Silphidae) of Wisconsin. Great Lakes Entomol. 38: 30-41.

Kriska, N. L., and D. K. Young. 2002. An annotated checklist of Wisconsin Scarabaeoidea (Coleoptera). Insecta Mundi 16: 31-48.

Lawrence, J. F. 1991. Endomychidae (Cucujoidea) (including Merophysiidae, Mycetaei-dae), pp. 482-485. In F. W. Stehr (ed.) Immature insects. Volume 2. Kendall Hunt, Dubuque, Iowa.

LeConte, J. L. 1954. Synopsis of the Endomychidae of the United States. Proc.Acad. Nat. Sci. Phila. 6: 357-360.

Leng, C. W. 1920. Catalogue of the Coleoptera of America, North of Mexico. John D. Sherman, Jr., Mount Vernon, New York.

Leschen, R. A. B., and C. E. Carlton. 1988. Immature stages of Endomychus biguttatus Say (Coleoptera: Endomychidae) with observations on the alimentary canal. J. Kans. Entomol. Soc. 61: 321-327.

Lisberg, A. E., and D. K. Young. 2003. An annotated checklist of Wisconsin Mordellidae (Coleoptera). Insecta Mundi 17: 195-202.

Majka, C. G. 2007. The Erotylidae and Endomychidae (Coleoptera: Cucujoidea) of the Maritime Provinces of Canada: New records, zoogeography, and observations on beetle-fungi relationships and forest health. Zootaxa 1546: 39-50.

Pakaluk, J. 1984. Natural history and evolution of Lycoperdina ferruginea (Coleoptera: Endomychidae) with descriptions of immature stages. Proc. Entomol. Soc. Wash. 86: 312-325.

Peck, S. B., and M. C. Thomas. 1998. A distributional checklist of the beetles (Coleoptera) of Florida. Arthropods of Florida and Neighboring Land Areas 16: 89.

Pellitteri, P., and G. M. Boush. 1983. Stored-product insect pests in feed mills in southern Wisconsin. Wisconsin Academy of Sciences, Arts and Letters 71: 103-112.

Price, M. B., and D. K. Young. 2006. An annotated checklist of Wisconsin sap and short-winged flower beetles (Coleoptera: Nitidulidae, Kateretidae). Insecta Mundi 20: 69-84.

Rauterberg, F. 1885. Coleoptera of Wisconsin. Proc. Nat. Hist. Soc.Wisc. 1885: 48- 62.Shockley, F.W., and J.V. McHugh. 2003. The Handsome Fungus Beetles (Coleoptera:

Endomychidae) of Great Smoky Mountains National Park. Available at: <http://department.caes.uga.edu/entomology/McHugh/GSMNP Endomychids.htm>

Skelley, P. E., and G. R. Burgess. 1995. Trochoideus desjardinsi Guérin found in Florida (Endomychidae: Trochoideinae). Coleopt. Bull. 49: 289-291.

Skelley, P. E., and R. A. B. Leschen. 2002. Family 92: Endomychidae, pp. 366-370. In R. H. Arnett, Jr., M. C. Thomas, P. E. Skelley, and J. H. Frank (eds.), American beetles, Vol. 2. CRC Press, Boca Raton FL.

Strohecker, H. F. 1986. Catalog of the Coleoptera of America North of Mexico. Family Endomychidae. U. S. Dep. Agric. Agric. Handb. Washington, DC.

Tomaszewska, K. W. 2000. Morphology, phylogeny and classification of adult Endomy-chidae (Coleoptera: Cucujoidea). Ann. Zool. Wars. 50: 449-558.

Wickham, H. F. 1894. The Endomychidae and Erotylidae of Ontario and Quebec. Can. Entomol. 26: 337-342.

Wickham, H. F. 1895. A list of Coleoptera from the southern shore of Lake Superior. Proc. Davenport Acad.Nat. Sci. 6: 125-169.


Yanega, D., and R. A. B. Leschen. 1994. Beetles associated with bee nests (Hymenoptera: Apidae) in Chiapas, Mexico, with descriptions of the immature stages of Vanonus balteatus Werner (Coleoptera: Aderidae, Endomychidae, Meloidae). Coleopt. Bull. 48: 355-360.

Young, D. K. 1984. Field records and observations of insects associated with cantharidin. Great Lakes Entomol. 17: 195-199.

Young, D. K. 1989. Notes on the bionomics of Xenomycetes morrisoni Horn (Coleoptera: Endomychidae), another cantharidin-orienting fungus beetle. Pan-Pac. Entomol. 65: 447-448.

Young, D. K. 1998. The fire-colored beetles of Wisconsin (Coleoptera:Pyrochroidae). Available at: <http://entomology.wisc.edu/~young/pyro/wipyroch.html>


1Department of Entomology, The Ohio State University, 1680 Ohio Agricultural Re-search and Development Center, Madison Avenue, Wooster, Ohio 44691, USA. 2Current Address: Warnell School of Forestry and Natural Resources, University of Georgia, Athens, Georgia 30602, USA. 3Department of Entomology, Michigan State University, 243 Natural Science Building, East Lansing, Michigan 48824, USA. *Corresponding author: (Phone: 706-542-4614; Fax: 706-542-8356; e-mail: [email protected]).

NEW REPORTS OF EXOTIC AND NATIVE AMBROSIA AND BARK BEETLE SPECIES (COLEOPTERA:

CURCULIONIDAE: SCOLYTINAE) FROM OHIODanielle M. Lightle1, Kamal J.K. Gandhi1, 2*, Anthony I. Cognato3, Bryson J. Mosley1,

David G. Nielsen1, and Daniel A. Herms1

ABSTRACTIn a 2007 survey of ambrosia and bark beetles (Coleoptera: Curculionidae:

Scolytinae) along a transect in northeastern Ohio, we collected six exotic and three native species not previously reported from the state. These species include the exotic ambrosia beetles Ambrosiodmus rubricollis (Eichhoff), Dryoxylon onoharaensum (Murayama), Euwallacea validus (Eichhoff), Xyleborus califor-nicus Wood, Xyleborus pelliculosus Eichhoff, and Xylosandrus crassiusculus (Motschulsky). The native ambrosia beetle Corthylus columbianus Hopkins, and the native bark beetles Dryocoetes autographus (Ratzeburg) and Hylastes tenuis Eichhoff are also reported from Ohio for the first time. Our study sug-gests a northward range expansion for five of the six exotic species including, X. crassiusculus, which is an important pest of nursery and orchard crops in the southeastern United States.

____________________

Exotic ambrosia and bark beetles (Coleoptera: Curculionidae: Scolytinae) cause significant ecological and economic damage to trees in forests, urban landscapes, and nurseries throughout North America (Kühnholz et al. 2001, Oliver and Mannion 2001). As global trade has increased in recent years, so has the number of exotic scolytine beetles detected and established in North America (Haack 2006). In response to the growing threat of invasion by exotic ambrosia and bark beetles, the United States Department of Agriculture Forest Service, Forest Health Protection branch (USDA Forest Service) established an Early Detection and Rapid Response (EDRR) program in 2001 (USDA For-est Service 2006). The main objectives of the EDRR program are as follows: 1) to monitor high-risk areas to detect and track recently introduced scolytine species; and 2) to respond rapidly to these new infestations to allow time for eradication programs. Ohio was among the 19 states surveyed as part of the EDRR program in 2007. In the 2007 survey in northeastern Ohio, we caught six exotic ambrosia beetles that had previously been unreported in literature. We also document three native ambrosia and bark beetle species not previously reported from Ohio.

From April through September 2007, scolytine beetles were sampled along a transect extending across five counties (Geauga, Holmes, Medina, Summit, and Wayne) in northeastern Ohio (Fig. 1). Nine trapping sites were monitored including six tree nurseries, an arboretum, a major interstate highway rest-area, and a semi-urban forested area. These sites were chosen because they


are considered high-risk for importation, movement, or establishment of exotic species, and/or are in proximity to ports-of-entry on Lake Erie. Scolytine beetles were sampled with 12-unit Lindgren funnel traps with a wet collection cup (Lindgren 1983). The collection cup contained 4-5 cm of non-toxic antifreeze to kill insects. Traps were baited with the following three semiochemicals: 1) ultra high-release ethanol (400 mg/day; chemical purity > 98%); 2) ultra high-release (+)-α-pinene (2 g/day; chemical purity 99%) and ultra high-release ethanol; and 3) exotic Ips bait consisting of (±)-ipsdienol (27 µg/day; chemical purity >95%), 2-methylbut-3-en-2-ol (30 mg/day; chemical purity >95%), and (±)- cis-verbenol (0.6 mg/day; 80% (-) enantiomer; chemical purity >95%) (Pherotech Interna-tional Inc.; Synergy Semiochemical Corp.). Lures were changed every 60 days. Three funnel traps were placed in each of the nine sites for a total of 27 traps for the study. The three traps were placed on a linear transect along forest or woodlot edges, and were separated by > 25 m to reduce inter-trap interactions. Traps were deployed from April to September and emptied every 14 days. All adult scolytine beetles caught in the study were identified to species by AIC and were cross-referenced with scolytine literature for new records (e.g., Wood 1982, Wood and Bright 1992, Bright and Skidmore 1997, 2002, Rabaglia et al. 2006). Voucher specimens are deposited at the Museum of Biological Diversity, The Ohio State University, Columbus, Ohio. Collection data for each specimen in the following section includes county, latitude and longitude, the bait used to capture that species, collection date, and total number of specimens collected (indicated parenthetically at the end of the record) (Table 1).

Figure 1. Map of Ohio showing location of detections of six exotic ambrosia beetles in 2007 Early Detection Rapid Response survey of northeastern Ohio. The symbols indi-cate locations of new species detected at each of the nine trapping sites (in some cases, there are multiple sites within a county).


ble

1. S

umm

ary

of n

ew s

tate

rec

ords

of s

coly

tine

beet

les

caug

ht a

s a

part

of t

he E

arly

Det

ectio

n Ra

pid

Resp

onse

(ED

RR) s

urve

y of

exo

tic

bark

bee

tles

in n

orth

east

ern

Ohi

o in

200

7.

Exo

tic/

B

ark/

Am

bros

ia

Num

ber

ofSc

olyt

ine

Bee

tle

Spec

ies

Nat

ive

Bee

tle

Adu

lts

Cau

ght

Ohi

o C

ount

ies

Ambr

osio

dmus

rubr

icol

lis (E

ichho

ff)

Exot

ic Am

bros

ia

5 La

ke, M

edin

a, W

ayne

Co

rthy

lus c

olum

bian

us H

opki

ns

Nat

ive

Ambr

osia

13

H

olm

es, L

ake,

Med

ina,

Sum

mit,

Way

ne

Dry

ocoe

tes a

utog

raph

us (R

atze

burg

) N

ativ

e Ba

rk

645

Gea

uga,

Lak

e, M

edin

a, S

umm

it, W

ayne

D

ryox

ylon

ono

hara

ensu

m (M

uray

ama)

Ex

otic

Ambr

osia

1

Lake

Eu

wal

lace

a va

lidus

(Eich

hoff)

Ex

otic

Ambr

osia

11

1 G

eaug

a, H

olm

es, L

ake,

Med

ina,

Sum

mit,

Way

ne

Hyl

aste

s ten

uis E

ichho

ff N

ativ

e Ba

rk

4 G

eaug

a, L

ake,

Sum

mit

Xyle

boru

s cal

iforn

icus

Woo

d Ex

otic

Ambr

osia

1

Lake

Xy

lebo

rus p

ellic

ulos

us E

ichho

ff Ex

otic

Ambr

osia

3

Sum

mit

Xylo

sand

rus c

rass

iusc

ulus

(Mot

schu

lsky

) Ex

otic

Ambr

osia

2

Way

ne


Exotic Scolytine BeetlesAmbrosiodmus rubricollis (Eichhoff)USA: Ohio, Lake Co., N 41°49’25” W 81°03’03”, N 41°36’37” W 81°18’57”,

(+)-α-pinene and ethanol, exotic Ips lure, 15.V.2007, 4.IX.2007 (2). Medina Co., N 41°04’09” W 81°44’10”, ethanol, 27.VI.2007 (2). Wayne Co., 40°46’53” W 81°54’57”, (+)-α-pinene and ethanol, 4.IX.2007 (1). Ambrosiodmus rubricollis is endemic to Asia and was first discovered in Maryland in 1968, and is now common throughout the southeastern United States. It has been reported from Alabama, Connecticut, Delaware, Florida, Louisiana, Maryland, Mississippi, Pennsylvania, South Carolina, Tennessee, and Virginia (Rabaglia et al. 2006). Common host species include Carya spp., Cornus spp., Prunus spp., and Quercus spp. (Wood 1982).

Dryoxylon onoharaensum (Murayama) USA: Ohio, Lake Co., N 41°36’37” W 81°18’57”, (+)-α-pinene and etha-

nol, 15.V.2007 (1). This exotic species also is distributed throughout the southeastern United States, and has been detected in Delaware, Florida, Georgia, Louisiana, Maryland, Mississippi, North Carolina, South Carolina, Tennessee, and Texas (Bright and Rabaglia 1999, Coyle et al. 2005). There is little known about the biology of D. onoharaensum. The known host spe-cies include Acer saccharum Marsh., Populus deltoides Bartr. ex Marsh, and Quercus spp. This species appears to be expanding its range, and threatens to become a pest of increasing economic significance in both forested and urban landscapes (Bright and Rabaglia 1999).

Euwallacea validus (Eichhoff)USA: Ohio, Geauga Co., N 41°28’31” W 81°20’28”, (+)-α-pinene and ethanol,

ethanol, exotic Ips lure, 15.V.2007- 20.VIII.2007 (7). Holmes Co., N 40°30’17” W 82°6’53”, (+)-α-pinene and ethanol, ethanol, 1-15.V.2007 (4). Lake Co., N 41°49’25” W 81°03’03”, N 41°49’07” W 81°02’18”, N 41°36’37” W 81°18’57”, (+)-α-pinene and ethanol, ethanol, exotic Ips lure, 15.V.2007-27.VI.2007 (30). Medina Co., N 41°04’09” W 81°44’10”, (+)-α-pinene and ethanol, ethanol, exotic Ips lure, 1.V.2007-7.VIII.2007 (18). Summit Co., N 41°14’03” W 81°31’35”, N 41°12’41” W 81°40’01”, (+)-α-pinene and ethanol, ethanol, exotic Ips lure, 1.V.2007-27.VI.2007 (49). Wayne Co., N 40°46’53” W 81°54’57”, ethanol, exotic Ips lure, 1.V.2007, 20.VIII.2007 (3). An exotic Asian species, E. validus was first reported in North America from New York in 1976. This species is now com-mon in the northeastern United States, and has been documented in Delaware, Louisiana, Maryland, New Jersey, New York, Pennsylvania, South Carolina, Virginia, and West Virginia (Rabaglia et al. 2006). Known host species include Abies spp., Picea spp., and Populus spp. (Wood 1982, Coyle et al. 2005). This species was caught in relatively high numbers in Ohio (111 individuals), and was present in all the six sampled counties (Figure 1).

Xyleborus californicus Wood USA: Ohio, Lake Co., N 41°49’07” W 81°02’18”, ethanol, 15.V.2007 (1).This palearctic species was first recorded in the western United States in

1944, and was not reported in the eastern United States until 2000 (Wood 1982, Vandenberg et al. 2000). It has been recorded from Alabama, Arkansas, California, Delaware, Florida, Kansas, Louisiana, Maryland, Mississippi, North Carolina, Oregon, South Carolina, Tennessee, Texas, Virginia, and Washington (Rabaglia et al. 2006). Hosts are unrecorded but individuals have been collected in Pinus taeda L. and Populus spp. stands (Fletchmann et al. 1999, Coyle et al. 2005).

Xyleborus pelliculosus EichhoffUSA: Ohio, Summit Co., N 40°45’15” W 82°21’42”, N 41°14’03” W 81°31’35”,

ethanol, 1.V.2007 (3). This Asian species was first reported in North America


in 1987 (Atkinson et al. 1990), and has been detected in Delaware, Maine, Maryland, Pennsylvania, Rhode Island, Tennessee, and Virginia (Rabaglia et al. 2006). There is no information about hosts or damage caused by this spe-cies in the United States, however hosts in Asia include Acer spp. and Quercus spp. (Haack 2006).

Xylosandrus crassiusculus (Motschulsky) (Granulate Ambrosia Beetle)

USA: Ohio, Wayne Co., N 40°46’53” W 81°54’57”, ethanol, 12-27.VI.2007 (2). The granulate ambrosia beetle is native to Asia and is especially prevalent in the southeastern United States. It has been recorded in Alabama, Delaware, Florida, Georgia, Hawaii, Indiana, Kansas, Kentucky, Louisiana, Maryland, Mississippi, North Carolina, Oregon, South Carolina, Tennessee, Texas, and Virginia (Rabaglia et al. 2006). Xylosandrus crassiusculus has dozens of hosts worldwide, including economically important plants such as Asimina triloba (L.) and Pryus calleryana Dcne., as well as Acer spp., Populus spp., and Quercus spp. (Horn and Horn 2006). The granulate ambrosia beetle is also known to attack healthy and newly transplanted trees, especially in nurseries (Solomon 1995, Oliver and Mannion 2001). The presence of this species in Ohio represents a major northward range-extension. As X. crassiusculus is one of the major pest in southeastern states (Kovach and Gorsuch 1985), it’s detection in Ohio is a case for concern for the eastern nursery and orchard stocks.

Native Scolytine BeetlesCorthylus columbianus Hopkins (Columbian Timber Beetle) USA: Ohio, Holmes Co., N 40°30’17” W 82°6’53”, (+)-α-pinene and etha-

nol, 15.V.2007 (1). Lake Co., N 41°49’25” W 81°03’03”, N 41°36’37” W 81°18’57”, (+)-α-pinene and ethanol, exotic Ips lure, 15.V.2007-10.VII.2007 (3). Medina Co., N 41°04’09” W 81°44’10”, (+)-α-pinene and ethanol, ethanol, exotic Ips lure, 15.V.2007- 7.VIII.2007 (4). Summit Co., N 41°12’41” W 81°40’01”, (+)-α-pinene and ethanol, ethanol, 24.VII.2007 – 7.VIII.2007 (3). Wayne Co., N 40°46’53” W 81°54’57”, ethanol, 29.V.2007, 24.VII.2007 (2). The Columbian timber beetle is a North American bark beetle that has been recorded from Delaware, Florida, Georgia, Indiana, Kansas, Maryland, Massachusetts, Missouri, New Jersey, New York, North Carolina, South Carolina, Tennessee, Vermont, Virginia, Washington DC, and West Virginia (Wood 1982, Rabaglia and Valenti 2003). The host species include deciduous trees such as Acer rubrum L., A. sacchari-num L., Castanea dentata (Marsh.) Borkh., Liriodendron tulipifera L., Platanus occidentalis L., Quercus alba L., and Ulmus spp. (Wood 1982). The Columbian timber beetle is economically important because it colonizes sapwood of healthy commercial trees, and can lower timber value by 25% (Solomon 1995).

Dryocoetes autographus (Ratzeburg)USA: Ohio, Geauga Co., N 41°28’31” W 81°20’28”, (+)-α-pinene and etha-

nol, ethanol, 15.V.2007-4.IX.2007 (323). Lake Co., N 41°49’25” W 81°03’03”, N 41°36’37” W 81°18’57”, N 41°49’07” W 81°02’18”, (+)-α-pinene and ethanol, ethanol, 15.V.2007-4.IX.2007 (223). Medina Co., N 41°04’09” W 81°44’10”, (+)-α-pinene and ethanol, ethanol, 29.V.2007-4.IX.2007 (27). Summit Co., N 41°14’03” W 81°31’35”, N 41°12’41” W 81°40’01”, (+)-α-pinene and ethanol, 29.V.2007-4.IX.2007 (21). Wayne Co., N 40°46’53” W 81°54’57”, (+)-α-pinene and ethanol, ethanol, 29.V.2007-4.IX.2007 (51). Dryocoetes autographus is native to North America with a transcontinental distribution. It has been recorded from Alaska, California, Colorado, Delaware, Idaho, Indiana, Maine, Maryland, Michigan, Minnesota, Montana, Nevada, New Hampshire, New Mexico, New York, North Carolina, Oregon, Pennsylvania, South Dakota, Tennessee, Utah, Virginia, Washington, West Virginia, Wisconsin, and Wyoming (Deyrup 1981, Wood 1982, Rabaglia and Valenti 2003). Due to its extensive distribution in the United States, it is surprising that D. autographus had not been previously


reported from Ohio. This species is typically found in the trunk and roots of dying trees and stems of fallen trees (Bright 1976, Wood 1982). Host species include Abies spp., Picea engelmannii Parry ex Engelm., P. glauca (Moench) Voss, Pinus contorta Dougl. ex. Loud., P. monticola Dougl. ex D. Don, P. strobus L., Pseudotsuga menziesii (Mirb.) Franco, and Tsuga heterophylla (Raf.) Sarg. (Wood 1982).

Hylastes tenuis Eichhoff USA: Ohio, Geauga Co., N 41°28’31” W 81°20’28”, (+)-α-pinene and etha-

nol, V.2007-VI.2007 (2). Lake Co., N 41°49’25” W 81°03’03”, (+)-α-pinene and ethanol, 15.V.2007 (1). Summit Co., N 41°14’03” W 81°31’35”, (+)-α-pinene and ethanol, 24.VIII.2007 (1). This North American bark beetle is especially abundant in the southern United States and has been recorded from Alabama, Arizona, Arkansas, California, Delaware, Washington DC, Florida, Georgia, Idaho, Indiana, Kentucky, Louisiana, Maryland, Massachusetts, Nevada, New Jersey, New Mexico, New York, North Carolina, Oregon, Pennsylvania, South Carolina, Tennessee, Texas, Utah, Virginia, and West Virginia (Wood 1982, Rabaglia and Valenti 2003). Hylastes tenuis is a rhizophagous species, coloniz-ing roots of various Pinus spp. throughout its range (Wood 1982).

In our study, five of the six exotic species (A. rubricollis, D. onoharaensum, X. californicus, X. crassiusculus, and X. pelliculosus) were previously known to occur primarily in southern states. These state records suggest a northward range expansion of these exotic species. It has come to our attention that some of the exotic and native species reported in this study also have been collected in Ohio by Robert A. Haack (USDA Forest Service), Robert J. Rabaglia (USDA Forest Service), and E. Richard Hoebeke (Cornell University) who retain the relevant collection records (all personal communication). To our knowledge, our study represents the first detection of X. crassiusculus in Ohio which is signifi-cant, as this beetle is a serious economic pest of nursery and orchard crops in the southeastern states (Kovach and Gorsuch 1985, Oliver and Mannion 2001).

ACKNOWLEDGMENTSWe thank the field and laboratory assistance provided by L. Aliaga, B.

Chambers, A. Claure, A. Font, D. Hartzler, and R. Neiswander (Department of Entomology, The Ohio State University), and R. Acciavatti (USDA Forest Service, Morgantown, WV, retired). V. Muilenburg (Department of Entomology, The Ohio State University) and two anonymous reviewers provided valuable comments on the manuscript. Funding for this project was provided by the USDA Forest Service, State and Private Forestry, Forest Health Protection, Early Detection and Rapid Response Project [# 07-DG-11420004-094 to DAH (The Ohio State University) and # 07-DG-11420004-182 to AIC (Michigan State University)], and by state and federal funds appropriated to the Ohio Agricul-tural Development and Research Center and The Ohio State University.

LITERATURE CITEDAtkinson, T. H., R. J. Rabaglia, and D. E. Bright. 1990. Newly detected exotic species

of Xyleborus (Coleoptera: Scolytidae) with a revised key to species in eastern North America. Can. Entomol. 122: 92-104.

Bright, D. E. 1976. The bark beetles of Canada and Alaska, Coleoptera: Scolytidae. The Insects and Arachnids of Canada, Part 2, Biosystematics Research Institute, Canada Department of Agriculture, Ottawa, Ontario, Canada. 241 pp.

Bright, D. E., and R. J. Rabaglia. 1999. Dryoxylon, a new genus for Xyleborus onoharaen-sis Murayama, recently established in the southeastern United States (Coleoptera: Scolytidae). Coleopt. Bull. 53: 333-337.


Bright, D. E., and R. E. Skidmore. 1997. A catalog of Scolytidae and Platypodidae (Coleoptera), Supplement 1 (1990-1994). NRC Research Press, Ottawa, Ontario, Canada, 368 pp.

Bright, D. E., and R. E. Skidmore. 2002. A catalog of Scolytidae and Platypodidae (Coleoptera), Supplement 2 (1995-1999). NRC Research Press, Ottawa, Ontario, Canada, 523 pp.

Coyle, D. R., D. C. Booth, and M. S. Wallace. 2005. Ambrosia beetle (Coleoptera: Scolyti-dae) species, flight, and attack on living eastern cottonwood trees. J. Econ. Entomol. 98: 2049- 2057.

Deyrup, M. 1981. Annotated list of Indiana Scolytidae (Coleoptera). Great Lakes En-tomol. 14: 1-9.

Fletchmann, C. A. H., M. J. Dalusky, and C. W. Berisford. 1999. Bark and ambrosia beetle (Coleoptera: Scolytidae) responses to volatiles from aging loblolly pine billets. Environ. Entomol. 28: 638- 648.

Haack, R. A. 2006. Exotic bark- and wood-boring Coleoptera in the United States: recent establishments and interceptions. Can. J. For. Res. 36: 269-288.

Horn, S., and G. N. Horn. 2006. New host record for the Asian ambrosia beetle, Xylo-sandrus crassiusculus (Motschulsky) (Coleoptera: Curculionidae). J. Entomol. Sci. 41: 90-91.

Kovach, J., and C. S. Gorsuch. 1985. Survey of ambrosia beetle species infesting South Carolina peach orchards and a taxonomic key for the most common species. J. Agric. Entomol. 2: 238-247.

Kühnholz, S., J. H. Borden, and A. Uzunovic. 2001. Secondary ambrosia beetles in ap-2001. Secondary ambrosia beetles in ap-parently healthy trees: adaptations, potential causes and suggested research. Int. Pest Man. Reviews 6: 209-219.

Lindgren, B. S. 1983. A multiple funnel trap for scolytid beetles (Coleoptera). Can. En-tomol. 115: 299-302.

Oliver, J. B., and C. M. Mannion. 2001. Ambrosia beetle (Coleoptera: Scolytidae) spe-cies attacking chestnut and captured in ethanol-baited traps in middle Tennessee. Environ. Entomol. 30: 909-918.

Rabaglia, R. J., S. A. Dole, and A. I. Cognato. 2006. Review of American Xyleborina (Co-leoptera: Curculionidae: Scolytinae) occurring North of Mexico, with an illustrated key. Ann. Entomol. Soc. Am. 99: 1034-1056.

Rabaglia, R. J., and M. A. Valenti. 2003. Annotated list of the bark and ambrosia beetles (Coleoptera: Scolytidae) of Delaware, with new distributional records. Proc. Entomol. Soc. Wash. 105: 312-319.

Solomon, J. D. 1995. Guide to insect borers in North American broadleaf trees and shrubs. Agriculture Handbook. 706. Washington, DC: U.S. Department of Agriculture, For-est Service. 735 pp.

USDA Forest Service. 2006. Early Detection and Rapid Response for Exotic Forest Pests. Available at: <http://www.na.fs.fed.us/ra/specialinitiatives/earlydetect/sib06_edrr.pdf>

Vandenberg, N. J., R. A. Rabaglia, and D. E. Bright. 2000. New records of two Xyleborus (Coleoptera: Scolytidae) in North America. Proc. Entomol. Soc. Wash. 102: 62-68.

Wood, S.L. 1982. The bark and ambrosia beetles of North and Central America (Co-leoptera: Scolytidae), a taxonomic monograph. Great Basin Naturalist Memoirs, No. 6. Brigham Young University, Provo. 1359 pp.

Wood, S. L., and D. E. Bright. 1992. A catalog of Scolytidae and Platypodidae (Coleoptera), Part 2: Taxonomic Index, Great Basin Naturalist Memoirs No. 13. Brigham Young University, Provo. 1553 pp.


DESIGNATION OF A NEOTYPE FOR MITCHELL’S SATYR, NEONYMPHA MITCHELLII (LEPIDOPTERA: NYMPHALIDAE)

Christopher A. Hamm1

The Mitchell’s satyr, Neonympha mitchellii French 1889 (Lepidoptera: Nymphalidae) was described as a new species based on a series of six males and four females collected by J. N. Mitchell from “Wakelee bog” in Cass County, Michigan (French 1889). French did not designate a holotype from this series. Much of French’s collection, and the original material included in the description, are thought to be lost (J. Shuey, M. Nielsen and J. Wilker, pers. comm.).

I did not find the syntype series of Neonympha mitchellii in potential re-positories including the collections of the American Museum of Natural History, Michigan State University, the University of Michigan, and the Field Museum of Natural History. Also, lepidopterists throughout the Great Lakes region did not have any knowledge of the whereabouts of the syntype series.

There is an exceptional need to designate a neotype for N. mitchellii be-cause its species boundaries have been questioned. Recently a new subspecies of N. mitchellii was described. Neonympha mitchellii francisi Parshall and Krall 1989 is thought to represent a southern regional morphological variant; the northern N. mitchellii were subsequently designated N. m. mitchellii (Parshall and Krall 1989). Aspects of this southern regional variation are present among individuals from northern populations. Also, the species boundary between N. mitchellii and Neonympha areolatus (Smith, 1797) (Lepidoptera: Nymphalidae) is blurred because the wing maculation characters used to distinguish species are continuous rather than discrete (Mather 1965, Scott 1986). Male genitalia characteristics were later used to distinguish N. mitchellii from N. areolatus (Parshall and Krall 1989), though these characters have subsequently been found to vary beyond the described limits (Goldstein et al. 2004, C. Hamm pers. obs.). Recent molecular work is also beginning to shed light on the relationships between N. m. mitchellii and N. m. francisi and a change in the taxonomic status of these subspecies may occur in the near future (Goldstein et al. 2004).

Neotype.- A male, vouchered in the Albert C. Cook Arthropod Collection at Michigan State University, bearing the following labels:

1) Cass Co., MI T5S R13W S30 “Wakelee Bog” Tamarack Swamp 4 July 1971 M.C. Nielson

2) EUPTYCHIA MITCHELLII French McDunn No. 101 Det M.C. Nielson

1Department of Entomology, Michigan State University. 204 Center for Integrated Plant Systems. East Lansing, MI 48824. (e-mail: [email protected]).


3) 000705 MI Lep Survey (barcode label)

I designate this male as the neotype of Neonympha mitchellii and, there-fore, of the nominate subspecies. This specimen does not vary in any distinguish-able way from French’s original description.

ACKNOWLEDGMENTSI would like to thank James Boone of the Field Museum of Natural History;

David Grimaldi and Suzanne Rab Greene of the American Museum of Natural History; Mark O’Brien of the University of Michigan; Anthony Cognato, Gary Parsons and Mogens Nielson of Michigan State University; Jay McPherson of Southern Illinois University; and James Wilker for their invaluable assistance during my research. This research was partially supported by a Michigan State University Plant Science Fellowship to Christopher A. Hamm.

LITERATURE CITEDFrench, G. H. 1889. A new species of Neonympha. Can. Entomol. 21: 25-27.Goldstein, P. Z., S. Hall, B. Hart, S. M. Roble, and J. Shuey. 2004. Evaluation of relation-

ships and conservation status within the Neonympa mitchellii complex (Lepidoptera: Nymphalidae). Report for the U.S. Fish and Wildlife Service.

Mather, B. 1965. Eupythchia areolata: Distribution and variation, with special reference to Mississippi (Satyridae). J. Lepidopt. Soc. 19: 139-160.

Parshall, D. K., and T. W. Krall. 1989. A new subspecies of Neonympha mitchellii (French) (Satyridae) from North Carolina. J. Lepidopt. Soc. 43: 114-119.

Scott, J. A. 1986. The butterflies of North America. Stanford Univ. Press. 583 pp. Smith, J. E. 1797. The natural history of the rarer lepidopterous insects of Georgia 2: 25.


NEW RECORD OF VANESSA VIRGINIENSIS (DRURY) (LEPIDOPTERA: NYMPHALIDAE) AS A HOST OF THYRATELES PROCAX (CRESSON) (HYMENOPTERA: ICHNEUMONIDAE)

David B. MacLean1 and John Luhman2

ABSTRACTOn 4 August, 2005 a male Thyrateles procax (Cresson) (Hymenoptera: Ich-

neumonidae) emerged from a pupa of Vanessa virginiensis (Drury) (Lepidoptera: Nymphalidae) which was contained within a screened cage in Cook County, MN. This is the first published record of Vanessa virginiensis as a host of Thyrateles procax. Adults of T. procax appear to be rare, as only two old records (both males) are known from Minnesota (University of Minnesota, Insect Collection)

____________________

This note reports the first published record of Vanessa virginiensis (Drury) (Lepidoptera: Nymphalidae) as a host of Thyrateles procax (Cresson) (Hymenoptera: Ichneumonidae). On 4 August 2005, a male ichneumon wasp, later determined by the second author to be T. procax emerged from a pupa of V. virginiensis which was contained within a screened cage in Cook County, MN. A late instar larva of V. virginiensis was discovered ca.10 days earlier wander-ing outdoors and placed in a cage along with three other V. virginiensis larvae that had been reared on Anaphalis margaritacea (L.) (Pearly Everlasting). A. margaritacea is a known larval host of V. virginiensis (Scott 1986) and is com-mon in Cook County, MN. Four larvae (including the one discovered outdoors away from its host plant) had pupated within the cage by late July. Two adult V. virginiensis emerged on 1 August 2005 and a third on 4 August. Additionally, on 4 August, a male T. procax emerged from the remaining V. virginiensis pupa. As all V. virginiensis larvae, except one, were reared from eggs collected on A. margaritacea and placed within a screened cage, we suspect that the late instar larva discovered wandering outdoors was most likely parasitized by T. procax. The adult wasp and the host pupa containing the last larval instar exuvium were pinned and placed in the first author’s collection. Adults of T. procax (both males) appear to be rare as only two other specimens of T. procax are known from Minnesota (University of Minnesota, Insect Collection).

Species of Thyrateles mainly parasitize larvae of Nymphalidae (species of Polygonia, Vanessa and possibly Oeneis) (Heinrich 1960). Peck (1964) lists V. virginiensis and Vanessa cardui (L.) as larval hosts of Thyrateles lugubrator (Gravenhorst) and Nymphalis antiopa (L.) as a host of T. procax. A search of the Zoological Record and Entomology Abstracts from 1980 to the present revealed no additional host records for T. procax.

LITERATURE CITEDHeinrich G.H. 1960. Synopsis of Nearctic Ichneumoninae Stenopneusticae with particular

reference to the northeastern region (Hymenoptera). Part III. Synopsis of the Ichneu-moni: genera Ichneumon and Thyrateles. Can. Entomol. Supplement 21. 367 pp.

1Professor Emeritus, Youngstown State University, current address: 76 Walter Rd. Grand Marais MN 55604. 2 Biological Control Scientist.


Peck, O. 1964. Synopsis of Nearctic Ichneumoninae Stenopneusticae with particular reference to the northeastern region (Hymenoptera). Part VIII. Addenda and corri-genda, host-parasite list and generic host index, Index to Ichneumonid names. Mem. Entomol. Soc. Canada No. 35: 889 - 925.

Scott, J. A. 1986. The butterflies of North America. Stanford Univ. Press. 583 pp.


FIRST OCCURRENCE OF HIPPODAMIA VARIEGATA (GOEZE) (COLEOPTERA: COCCINELLIDAE) IN OHIO

Daniel M. Pavuk1, Alan Sundermeier2, Sadira Stelzer1, Andrea M. Wadsworth1, Danielle M. Keeler1, Melanie L. Bergolc1, and Laura Hughes-Williams1

Ladybird beetles, or coccinellids (Coleoptera: Coccinellidae), are significant arthropod predators in a variety of terrestrial ecosystems. Numerous classical bio-logical control projects undertaken over the last 120 years in North America have involved importation of exotic ladybird beetle species for the control of invasive insect species in annual and perennial agricultural production systems. Some examples of coccinellid species imported into North America in the last 40-50 years include the seven-spotted ladybird, Coccinella septempunctata L., the variegated ladybird, Hippodamia variegata (Goeze), and the multi-colored Asian ladybird, Harmonia axyridis (Pallas). Some of these imported coccinellids have established quite suc-cessfully in many habitats of the continent, and in at least certain situations, have demonstrated significant prey population regulation capacity.

As part of a study conducted in 2006 and 2007 to document arthropod preda-tor communities in edge habitats associated with soybean agroecosystems, we were interested in determining if the nonnative coccinellid, H. variegata, was present in Ohio soybean agroecosystems. Much effort has been directed at documenting predatory coccinellids associated with soybean agroecosystems since the invasion and establishment of the soybean aphid, Aphis glycines Matsumura (Hemiptera: Aphididae), in North America. This alien aphid has become a major pest of soybeans in the North Central Region since being first observed in 2000 in Wisconsin (Alle-man et al. 2002). Gardiner and Parsons (2005) detected H. variegata in Michigan in 2005, so we wished to know if this coccinellid had moved into Ohio.

Nonnative and native coccinellid species captured by sweep net are listed in Tables 1 and 2, respectively. In both years, H. axyridis was a dominant species in the collections. Larger numbers of coccinellids were collected in 2007 than in 2006, and a greater number of coccinellid species was collected in 2007 than in 2006. Interestingly, H. variegata was not collected in soybean fields in 2006, but was the second most common coccinellid species sampled in 2007. This represents the first published record for H. variegata in Ohio. It appears that H. variegata is spreading eastward from its original point of detection in Montreal in 1984 (Gordon 1987).

1Department of Biological Sciences, Bowling Green State University, Bowling Green, OH 43403-0208. 2The Ohio State University, Wood County Extension, 639 Dunbridge Rd., Suite 1, Bowling Green, OH 43402.

Table 1. Nonnative Coccinellid Species Associated with Soybean Agroecosystems.

Year Coccinella septempunctata Hippodamia variegata Harmonia axyridis

2006 4 none collected 202007 30 26 21

Table 2. Native Coccinellid Species Associated with Soybean Agroecosystems.

Year Hippodamia Cycloneda Coleomegilla Brachiacantha Hippodamia parenthesis munda maculata ursina convergens lengi

2006 none collected 1 2 none collected none collected2007 6 6 4 4 2


ACKNOWLEDGMENTSWe wish to thank numerous soybean growers in the northwest Ohio region

who allowed us to sample arthropods in their soybean fields. We also thank Luke Sundermeier for his assistance in sampling arthropod populations. This research was supported by a grant from the Warner Endowment Fund for Sustainable Agriculture at The Ohio State University awarded to A. Sundermeier and D. M. Pavuk and research funds and support from the Department of Biological Sciences, Bowling Green State University.

LITERATURE CITEDAlleman, R. J., C. R. Grau, and D. B. Hogg. 2002. Soybean aphid host range and virus-

transmission efficiency. Proc. Wisc. Fertilizer Aglime Pest Manage. Conf. Available at: <http://www.soils.wisc.edu/extension/FAPM/2002.php>

Gardiner, M. M., and G. L. Parsons. 2005. Hippodamia variegata (Goeze) (Coleoptera:Coccinellidae) detected in Michigan soybean fields. Great Lakes Ento-mol. 38: 164-169.

Gordon, R. D. 1987. The first North American records of Hippodamia variegata (Goeze)(Coleoptera: Coccinellidae). J. N. Y. Entomol. Soc. 95: 307-309.

Documents

THE MICHIGAN ENTOMOLOGICAL SOCIETY ...2007 THE GREAT LAKES ENTOMOLOGIST 103 1Lepidoptera Alert a.k.a. Lepalert, 2806 Linwood Avenue, Royal Oak, MI 48073-3023. (e-mail: [email protected])