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MICHIGAN STATE UN NE

1 : :1 111.111:11111111111111:

31293 00606 0515
LIBRARY
Michigan State
University

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

This is to certify that the

dissertation entitled
iRAP-IESIIIG BIOLOGY OF
PASSALOECIIS CIISPIDAIIIS SHIIII (IIYIIEIOPIERA: SPECIDAE)

AID SYHPAIRIC SPECIES
presentedby

 

John Morris Fricke

has been accepted towards fulfillment
of the requirements for

"1.0. degreein Entogology

RMMQ Agra 4

Major professor

Date 16 Februarl 1990

MS U i: an Affirmative Acn'on/ Equal Opportunity Institution 0-12771

 

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TO AVOID FINES return on or before one due.

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MSU Is An Affirmdive Action/Equal Opportunity Institmion

 

 

 

 

 

 

 

TRAP-NESTING BIOLOGY OF

EASSALQEQQS QHSELDAIQS SMITH (HYMENOPTERA: SPHECIDAE)
AND SYMPATRIC SPECIES

BY

John Morris Fricke

A DISSERTATION

Submitted to
Michigan State University
in partial fufiiiment of the requirements
for the degree of

DOCTOR OF PHILOSOPHY

Department of Entomology

1990

Trip-nee
CIIEEO out
Waugh 19:
Partitioning
Dore dime:
(3a?) and E
‘ 6.4 m,

ABSTRACT
TRAP-NESTING BIOLOGY OF
EASSALQEQQS CQSBLDAIQS SMITH (HYMENOPTERA: SPHECIDAE)
AND SYMPATRIC SPECIES
BY

John Morris Fricke

Trap-nesting studies of Eaaaalgggus (Hymenoptera: Sphecidae) were
carried out at Concordia College, Ann Arbor, Michigan, from 1984
through 1987. Variations in nest architecture and resource
partitioning among sympatric Eassalgggua spp. were studied. Trap nest
bore diameter selections among W spp. were: 2. annulus]:
(Say) and E. angglatus Vincent, 1.6 - 2.4 mm; B. guspigatua Smith, 2.0
- 6.4 um, and E. W Dahlbom, 1.6 - 3.6 mm. B. W
preferred trap nests on Rings and E. angglatua preferred trap nests on
Jgglana. E. angglatua nests were most frequent at heights of 3 to 9
m; E. W nests were most frequent below 3 m. mm: mm
(Fabricius) (Chrysididae) parasitized nests of E. annulatus, E,
arenlatuss E. suspidatus. and E. mpnillccnnis: Qmaius, inidescens
(Norton) parasitized E. annuiatua; and Inignnxsls dozing Gribodo
parasitized E. gusgigatua. Egmgnia Luggagiga (Cresson)
(Ichneumonidae) parasitized B. W and Bangui; AIMEE:
parasitized E. W, B. W,» and B. W. Mean

length of provisioned cells decreased as bore diameter increased, and
volumes of provisioned cells increased with increasing bore diameter.

Increased length of provisioned cells was associated with higher

   
   

parasite activity
itapnest sore ope:
mm spp.
«creases in bass
cell lengths. A 1
trap nests with

I“ 1987. provisl
thmg‘ AUgh'st E
am 9618 aphids

the average nun:

parasite activity but not with wasp senescence. Orientation of
trapnest bore openings did not influence frequency of trap nest use by
Eassalgggus spp. Decreased bore depth resulted in significant
decreases in basal cell lengths and slight decreases in provisioned
cell lengths. A signficant decrease in number of provisioned cells in
trap nests with 60 mm bore depths was noted only for E. gugpiggtus.
In 1987, provisioning activity of E, ggspiggtua extended from May 29
through August 6. Eighty trap nests contained 281 provisioned cells
and 9618 aphids. The average number of aphids per cell was 34.2 and
the average number of cells provisioned per day per trap nest was .73.
B. We used mm 89-. W 89.. W 8:».
Magnggipnum gypngnnlgg (Thomas), Myzug sp., Myzua gggggi (Fabricius),
Minus monagggg (Davis), and filtgpiumyaggnag Fabricius. The number of

aphids provisioned per cell was significantly different among three
species of aphids and was inversely related to aphid size. Resin
gathering activities of E. M were also observed and

described.

DEDICATION

This work is dedicated to Henry and Rose Muschick

for their love and encouragement.

iv

he author
institutions and
development of 1
which granted
38i1982; Aid
apport curing
Gary Simmons,
(“Ci-39m State
Hithigan; my \
long-miter ing
p“(leasing a
invertebrate
iianya StoetZE

Roiand L

Fis

ACKNOWLEDGEMENTS

The author expresses his appreciation to the following
institutions and individuals that contributed to the initiation and
development of this study: Concorida College, Ann Arbor, Michigan,
which granted a sabbatical study leave for the academic year
1981-1982: Aid Associaton for Lutherans, which provided financial
support during the study leave; Drs. Edward Grafius, Frederick Stehr,
Gary Simmons, and Steve Stephenson, Guidance Committee members
(Michigan State Univeristy); Dr. John Witter, advisor (University of
Michigan; my wife Karen, and children Jeff and Jennifer for their
long-suffering patience and word processing; Maryann Oester, word
processing and deciphering of hand written manuscripts: the
invertebrate zoology class of 1986 for trap nest construction; Dr.
Hanya Stoetzel, USDA, for the identification of prey: and finally, Dr.

Roland L. Fischer. guidance comnittee chairman, and mentor.

LIST OF TABLES
LIST OF FIGURES
IKEQEKKISN
$323038 AND RX
RESJL'TS AND DI

Nxo Dian
E§§§aioec

Trap Res
Parasite
Provisi
Factors
Bore Di
Bore E

Berg 1
Trap

Bor e

lnfl
TYQ‘

bob

the
3%AP“.

TABLE OF CONTENTS

LIST OF TABLES .........................
LIST OF FIGURES .........................
INTRODUCTION ..........................
METHODS AND MATERIALS ......................
RESULTS AND DISCUSSION .....................
Bore Diameter Preferences Among Sympatric
Easaalgggus spp ....... . ..... . .........
Trap Nest Height Selection in Raggalggggg spp ........
Parasites of Eaasalgggua spp. . . . . . . . ....... .
Provisioned Trap Nest Architecture .............
Factors Influencing Cell Length and Volume ...... . . .
Bore Depth and Cell Architecture ..............
Bore Depth and Number of Provisioned Cells .........
Bore Opening Orientation and Frequency of
Trap Nest Use . . . . . . . . . . . . . . . ........
Bore Diameters and Number of Provisioned Cells .......

Influence of Station Species on Frequency of
Trap Nest Use . . . . . . . . . . . . . . . . .......

Aphid Provisioning by Easaglgggu§,gu§gigatug ........

Economics of Cell Partitions and Closures .........

SUNNARY AND CONCLUSIONS .....................

vi

xii

17

17
26

64

71

87

AWDDHES . ..
APPENDIX i.
APPEIDIX 1.1

 

 

 

“TDNHX 2.

APPENDIX 3.

Intros
Biolog
Aphio
Key t
Hich?
List
Lite
kPttmnx

APPENDICES
APPEN
APPEN

APPEN

APPEN

vii
TABLE OF CONTENTS (cont’d)

........................... 9O
DIX 1. Record of Deposition of Voucher Specimens . . 9O
DIX 1.1 Voucher Specimen Data ............ 91

APPENDIX 2. Preliminary Surveys and Selection
of Study Area ................ 103
DIX 3. Ragaalgggus of Michigan
(Hymenoptera: Sphecidae) ........... 113
Introduction ....... . ............. 113
Biology of North American Eaasalgggua ......... 115
Aphid Prey of Pasaalgggua . . . . . . . . . . . . . . . 116
Key to Nichgian Easaalggcus .............. 117
Michigan Baasalgggua . . ............... 121
List of Figures (Appendix 3) ............. 136
Literature Cited . . . . . . . .......... . . 142
DIX 4. Data Tables from Trap-nesting
Studies of Baggalgggng, 1984-1987 ....... 144
DIX 5. Proposed Future Investigations . . ...... 163

APPEN

LITERATURE CITED . . . . . . . ............. . . . . 165

Table 1. Plant
Table 2. Bore l
1984-
IEDIE 3. E g
1984-
Table 4. Trap
Baa:
Table 5, DIsi
Ea:
Table 6 Pas
anc
Die
E3“.
Tam" 8- Di
5;:
Table 9. II
Table IO. 5
1

TabIe 13

 

Table

Table

Table

Table

Table

Table

Table

Table

Table

Table

Table

Table

Table

10.

11.

12.

13.

LIST OF TABLES

Plant cover of trap-nesting study area . . . .....

Bore diameter selections for five Eassalgegus spp. ,
1984-1986.. . ...... . . ....... . .

E. gusgidatus bore diameter selections,

1984-1987 ........... . ...........

Trap nest heights and frequency of use by four

Eassalgggua spp., 1984-1987 ..............

Distribution of Qma1u§,agngu§ (Fabricius) among
Passalggcus gusgigatus trap nest stations, 1984

Pasasitism of Eassalgg§u§_gu§21gatg§ trap nests
and cells by angina agngua (Fabricius) by Bore

Diameter, 1984 . . . . . . . . . . . . . . . . . . . .

Distribution of angina agngua and Bangui; among

Eassalgecua trap nests, 1987. . . . . ........

Distribution of Parasitism among Eassalgegua

guspigatus trap nests, 1987.. . . . . .......

Nean provisioned cell lengths for three Easaalgggns

spp. from pooled samples, 1984-1987 ........ . .

Simple linear regression of bore diameters and mean

cell lengths for three Eassalgggug spp. .......

Bore diameters and provisoned cell volumes for three

W app. 0 o e e e e e o e e o e e e o o e e 0

An analysis of number of aphids [uyzu§,mgnangag
(Davis)] provisioned per cell, cell lengths and cell
volumes in 3.2 mm bore and 4.0 mm bore trap nests

provisioned by E. ggggigatus, 1987. . . . . . . . . .
Seasonal distribution of Eaasalgggua ggsgigatga cells

of extra-ordinary length ..........

viii

16

21

23

28

3O

31

32

35

4O

41

4a

45

50

Table 14. Cell 1:
nests ;

Table 15. Cell l1
nests 1
E. m

 

Table 16. Hean o
and tw
Basia;

Table 17. Mean t
1986.
Table 18. Hean
3P?”
Table 19. iiean
3%.
Table 21L Nean
SPF.
Tdble 21. BC?!
by 1
Table 22- Fre

Tame 23.
We 24.

We 25.

We 27,

Table

Table

Table

Table

Table

Table

Table

Table

Table

Table

Table

Table

Table
Table

14.

15.

16.

17.

18.

19.

20.

21.

22.

23.

24.

25.

26.
2?.

ix
LIST OF TABLES (cont’d)

Cell length data from 1984 Eassaigegus guspidatus
nests parasitized by 0,. agnegs. . . . . . . . . 51

Cell length data for cells from non-parasitized trap
nests and parasitized trap nests provisioned by

2, guanigatns, 1984. . . . . . . . . . . . . . . . . . 53

Mean basal cell lengths from eight bore diameter classes
and two bore depth classes. Pooled data from

Easaalgggus trap nests, 1986. ............ 56
Mean basal cell lengths for Eassalgggus spp. ,
19860 0 O O O O O O O O O O O O O O 0 O O O 57

Mean lengths of provisioned cells of Eassalgegus
spp.,1986 ..... ..............58

Mean vestibular cell lengths for Easaalgegus
SP? . 1986. . . . . . . . . . . . ....... 59

Mean number of cells per trap nest for Easaaiagggus
spp., 1986 . . . . . . . . . . . . . . . . . . . . . . 61

Bore opening orientation and frequency of use
by Passaigggus spp.. 1984 and 1986. . . . . . . . . . 63

Frequency of E, guanigatus trap nest cells from 120
mm bore trap nests from 1984, 1985, and 1987. . . . . 65

Station selection by three Easaalgggus spp., 1986. . . 67

Frequency of trap nest use at secondary site by
all trapnesting wasps and bees, and by

Eassaigggus spp.. 1986. ............... 69
Station species frequency and frequency of trap

nest use by Eassalgggus ggsgidatua, 1987. . . . . . . 70
Aphids provisioned by E. gnaglgatua, 1987 ....... 75

Families of Mymenoptera collected at Concordia College,
Ann Arbor, Michigan, during the Summers of 1982 and 1983
using a malaise trap of Towne’s design. . . . . . . . 104

Table 28.

Table 29.
Table 30.

Table 31.
Table 32.
Table 33,

Tania 34.

Male 35.
Male 35_
Table 37_

We as.

Table 39.

SUD
at 1
Sum

m

Fre<

Rea

Table

Table
Table
Table

Table

Table

Table

Table

Table

Table

Table

Table

28.

29.
30.
31.

32.

33.

34.

35.

36.

37.

38.

39.

X

LIST OF TABLES (cont’d)

Sub-families and tribes of the Sphecidae collected

at Concordia College, Ann Arbor, Michigan, during the
Summers of 1982 and 1983 using a malaise trap of
Towne’s design . . . . . . . . . . . . . . . . . .

Frequency of Use By Nest Material Type ........

Rearing data for W maidens.

Bore diameters and lengths of Eagsalgegus
EHSELQBLHS cells. 1984.

Frequency of trap nest use and number of adults reared

out for four Eassalgggus spp., 1985. . . .

Frequency of trap nest use by four Eassalgegus
spp. , 1986 ..... . ....... . . .

1984.

Basal cell lengths from eight bore diameter classes
and two bore depth classes. Data compiled from four

W spp.. 1986 .

mm: mm: provisioned cell
lengths, 1986 ......

We mm: provisioned cell
lengths, 1984-1987.

OOOOOOOOOOOOOOO

W mum; provisioned cell
lengths, 1984-1986. . .

Lengths of Raggaigeggs provisioned cells from eight
bore diameter classes and two bore depth classes. Data
were combined from four Eassalgggus spp., 1986. . . .

Provisioned cell lengths for four Eassalgggus spp.,
1986. Cell lengths are compared on basis of bore
diameter and bore depth ................

105
111

144

145

146

146

147

148

149

151

152

153

fuflelo.

nanny

Tuflelz,

Infle 43.

7 ‘le 44.

Tune 45.

Provls
1986 u

Numbe'
bore .
Combi
Freq:
Rafiq

Vest
Clas
tou

Table

Table

Table

Table

Table

Table

40.

41.

42.

43.

44.

45.

xi

. LIST OF TABLES (cont’d)

Provisioned cell lengths for four Easaaigegus spp.,
1986 with the application of the 10% exclusion rule. . 154

Number of Eassalgggus provisioned cells from eight
bore diameter classes and two bore depth classes.
Combined data from four Eassalgegus spp., 1986. . . . 155

Frequencies of trap nest height selection by four

Eassalgegus spp., 1986. .......... . . . . . 155

Vestibular cell lengths (VCL) from eight bore diameter
classes and two bore depth classes. Data compiled from

four Eassalgggga spp., 1986. . . ..... . . . . . . 156
Cell Data on Aphids Provisioned by Eassaigegus
guagiggtug, 1987. .................. 157

Trap Nest Data on Aphids Provisioned by Eassaigegus
gnsgigatu§,1987. . . . . . . . . . . . . . . . 160

Figure l. Cons:
l. P.‘
remo:
IOUte
with

Figlirl 2. A 3 .

“We 3. Dis-t:

E19“? 4- ExpeI

“9‘" 5. Nest
Pm" 5- Scat
FlWe 7.

Figure B.

 

 

“er 9.

“wt-e 10.

“We 11

 

Figure

Figure

Figure

Figure

Figure

Figure

Figure

Figure

Figure

Figure

Figure

10.

11.

LIST OF FIGURES

Construction of pre-split trap nests.

A. Pine trap nest block; B. A 6.4 mm section

removed from side of trap nest; C. 60 degree channel
routed in trap nest face; D. Pre-split trap nest

with bore drilled out ................. 10
A 3 x 3 bundle of trap nests ready for distribution

into the field.

Distribution of bore openings in 3 x 3 and 4 x 5
trap nest bundles illustrating the opposite ends of
representative bundles.

Expected and observed fequencies of bore diameter

selection by Ea§§a1gg§ua gu§21datg§, 1984-1987. . . .

Nest architecture illustrating basal cell, provisioned
cells, vestibular cell, preliminary plug, resin
partitions and closure. . . . . . .

24

Scatter plot of number of aphids provisioned per cell
and cell volumes <mm3) for E. gu§21ga1g§ 3.2 mm

bore trap nests, 1987. 47

Scatter plot of number of aphids provisioned per cell
and cell volumes (mm3) for E. ggsg1g§1g§ 4.0 mm

bore trap nests, 1987 ................. 48
Frequency of ringed and closed trap nests during

10- -day intervals beginning May 29 and ending

August 6, 1987. . . ........
Cummulative frequencies of ringed and closed trap
nests during 10-day intervals beginning May 29

and ending August 6, 1987 ............... 74

Seasonal changes in prey selection by

B. W. Sumner 1987.

Number of aphids provisioned by B. gu§21dagu§
during 5-day intervals beginning May 29 and
ending August 6, 1987 .................

78

xii

Rome 12.

Figure 13.

P1 re 14.

Figure 15.

Figure 16.

xiii

LIST OF FIGURES (cont’d)

Figure 12. Cummulative number of aphids provisioned by
B. §n§21datus during 5-day intervals

beginning May 29 and ending August 6, 1987. ..... 79
Figure 13. Provisioned cell production rates for

E. QMSELQBLHS ..................... 82
Figure 14. Thickness of resin partitions and closures

in E. gu§21ga1u§ trap nests .............. 86

Figure 15. Schematic map of Concordia College, Ann Arbor,
Michigan, indicating trap-nesting sites for
Summer 1982 ..................... 106

Figure 16. Schematic map of Concordia College, Ann Arbor,
Michigan, indicating trap-nesting sites for
Summer 1983 ..................... 110

 

lian and the
inter-rel at i onsh i
lnetnnentai in
recreational pur
chains as bio
tWildcats, re
.cuts for pg”
“Never, 1
“Net; have C
”than, m
“n “006. m
“MOI. of Del
A: Moan
m'rpl‘izes.
for m“? e)
xaie meta
distribution

INTRODUCTION

Man and the insects have a long history of positive and negative
inter-relationships. Many insects directly or indirectly are
instrumental in providing food, fiber, shelter, and enJoyabie
recreational pursuits. Ecologically, insects have their place in food
chains as biological concentrators (herbivores and carnivores),
decaposers, remcers, or as benefactors to plants requiring biotic
agents for pollination.

However, when man manages ecosystems for his own purposes,
insects have opportunity to exploit concentrated resources and have
negatively influenced man’s obJectives. Any product of interest to
man (food, fiber, shelter, health, and recreation) is also a resource
capable of being used by insects for their own purposes.

As hulan activity shifted from subsistance levels to canerciai
enterprizes, resources were concentrated, increasing the opportunity
for their exploitation by insects. New technologies enabled larger
scale operations and increased travel provided opportunity for
distribution of potential insect pests. Energy investment,
prodnctivity, and profits increased. Monoculture replaced diverse
ecosystems: and accidental introdnctions of potential pests into rich
resource bases, lacking appropriate biological controls, often

resulted in staggering losses.

Bill
the agi‘
mound
nicotine
naterial
and pr!
Progress
receive:
mi in
sicknes:
Vectors
priliti‘
(Helm
Cultura

'ih
mrlng
“isect.
Comet.
“Shem
'CODtr;
Given
“tau.

p,
prii'iCi

Co

The re

Efforts to cmbat competitors for products of interest predate
the agricultural revolution. Early control techniques included
capounds of sulfur, arsenic, and copper; plant derivatives such as
nicotine, pyrethrins, and rotenone: and other naturally derived
materials such as chalk, wood ash, oil sprays, bitumen sticky bands,
and predatory ant colonies. As the agricultural revolution
progressed, pest control measures became more scientifically based and
received a tremendous imetus from the field of medical entomology as
many tropical diseases including Texas cattle fever, African sleeping
sickness, malaria, and yellow fever were shown to have arthropod
vectors. A mlti-faceted approach, which midnt be considered
primitive integrated pest management, emerged and included the
development of resistant varieties, and the imlimentation of
cultural, biological, and chemical controls.

The search for improved chemical control received great impetus
daring world war II with the realization that dangers of tropical,
insect-vectored diseases were surely as great as those of armed
coubat. Subsequent discovery of the insecticidal properties of DDT
ushered in a new age of optimism regarding insect management, and
'control'I of pests was replaced with the attitude of eradication.
Given this prospect, old pest management practices including crop
rotation, sanitation, and encouragnent of natural enemies declined.

Pesticide use accelerated, without regard for ecological
principles or consequences, and individual users applied chemical
controls even in situations where pests were not economically harmful.

The realization that mthing was amiss developed gramaily. with,

Hosxin
hay (2
attent

01 pe

targe‘
repro
popul
iollo
natur
Femov
non-t
dire:

of ti

and

reap
ette
test

Prof

(19:
Of 1
Y'iltl
to

€001

ell

iioskins, and Fuilmer (1948) reported that dairy cows fed low residue
hay (2 ppm DDT) produced butterfat with 65 ppm DDT, but little public
attention was focused on the persistance and biological magnification
of pesticides until the 1962 publication of Rachel Carson’s 511“;
m. Subsequently, four significant observations were made: 1)
target populations, especially those with short life cycles and hidl
reproductive capacities developed resistance to pesticides; 2) target
populations frequently rebounded to even higher population levels
following pesticide applications, often die to the elimination of
natural controls: 3) secondary pests became maJor pests due to the
removal of natural control agents; and 4) significant numbers of
non-target organisils were succunbing to pesticides as a result of
direct application, food chain magnification of ingested pesticides,
or the long term accumulation of hard pesticides with long half-lives.

In response to the development of resistance, application rates
and frequency of applications increased. The pesticide inmstry
responded with the development of alternative pesticides: but
effective life spans of pesticides decreased, and development and
testing costs contributed to increased cost of control, decreasing
profit margins.

aaith and Micheibacher (1949) and Smith, Anderson, and Reynolds
(1950) suggested that pest management should involve field evaluation
of pest populations and their natural enemies. This is associated
with the realization that pest populations can exist at levels too low
to Justify the cost of certain management practices. Gradually,
control systems have been developed which involve a holistic ecosystem

approach called Integrated Pest Management (1PM).

IP11
of crop
condit 1:
Classic:
scale bl
identif:

it
general
increas
precipi
them.
consequ
intereg
funding
incorpg
One an
inning

P
these
incluo
hey in
genera
beetle
a... .
”5393
term

cWill

1PM system success requires an accurate knowledge of the biology
of crops, pests, natural controls, and the effects of climatic
conditions on the population growth of each of these couponents.
Classical success stories, such as the control of cottony cushion
scale by the vedalia beetle, illustrate that conprehensive studies can
identify effective agents of biological control.

It is interesting to note that research efforts to learn the
general biology of insects and their biological control declined with
increased pesticide production and testing. The very actions that
precipitated greater need for such studies were mitigating against
them. The basic biology and potential use of natural enemies has
consequently been limited to those systems involving prodncts of
interest with the greatest econanic impact and hidnest potential
funding levels. Many predator-prey relationships await study and
incorporation into the knowledge base of pure and applied entomology.
One such relationship is that between the genus W and
Aphids.

Field studies of W show the correctness of describing
these wasps as aphid hunters. However, these was have not been
included in lists of aphid predators or parasites (parasitoids). This
may be the to the peculiar biology of the aphid hunting wasps in
general. These wasps provision nests located in the ground, twigs,
beetle borings, reeds, grass culls, and bark. Nests may be located
sane distance from an aphid colony and time spent by provisioning
wasps at these colonies may be minimal. This contrasts with the long
term close contact with an aphid colony by predaceous larvae of

coccinellids, chrysopids, syrphids. or hymenopterous parasitoids.

Di)
in man-
natural
(1975)
lungram
diiilcu
long 9:
in rear
greenho

Pr
serenoi
2- m:
dild Vll
1978).
d1Ainetl
of 3,2
Mat c

the a!

Dixon (1973) notes the success of biological controls on aphids
in man-managed ecosystems but indicates there is no evidence that
natural enemies regulate aphid populations. Corbett and Backhouse
(1975) suggest that aculeate Hymenoptera might be useful in 1PM
programs if mre were known of their biology. They recognize
difficulties (high temperature thresholds for activity, low fecundity,
long generation time, slow response to prey density, high investment
in rearing time, and availability of nest sites) but suggest that in
greenhouses management of these wasps could prove successful.

Previous work on Eas§a1gggu§ has been incidental and
serendipitous. The most comprehensive studies have been reported on
2.,gugp1gagus Smith (Fye, 1965b: Krombein, 1956, 1958, 1963, and 19673
and Vincent, 1978) and 2. W111: Dahiban (Fye, 1965b; Vincent.
1978). However, the scope of these studies was limited. Bore
diameters used in these trap-nesting studies usually had lower limits
of 3.2 m and rather large scale increments of 1.6 m. Bore diameters
most canoniy used have been 3.2, 4.8, 6.4, and 8.0 m. Considering
the snail size of W spp., snalier bore diameters are more
appropriate for determining bore diameter preferences. Data from
large bore trap nests suggest an inverse relationship between bore
diameter and cell length or cell volume. The effectiveness of smaller
bore diameters needs to be examined.

No biological studies have been reported on sympatric
Eassa1gggus. If several Eassa1gggus spp. are present in a community,
how are resources (aphids, nesting sites. closure materials, etc.)

partitioned? An abundance of resources midst make direct competition

sugge!

excius

599. l

have
activi
inlll'

and F

t

reporl

in Jul

Archi:

Stiecl

or interference of no consequence, but competition theory would
suggest that any advantage to one species would result in ultimate
exclusion of others, if the same resoures were required (Cause, 1934,
1935; Park, 1948). Can a partitioning of resources by W
spp. be demonstrated?

Corbet and Backhouse (1975) suggested that Eassa1gegns females
have a hunting life of 50 days, but only one report confirms an
activity period near that duration. Kronbein (1963) reported 2.
W as mltivoltine and active from May 20 through October 13
and E. W active 39 days (June 1 — July 9). Fye (1965b)
reported on E, mgn11ggnn1s_with a spring generation provisioning nests
in June and a summer generation provisioning nests in August.

Details on the nature of trap-nesting sites (exposures, heights,
station species, and plant caununities) have not been included in
previous studies. Such information would be helpful in determining
the factors influencing the local distribution of sympatric
W. The research reported below was carried out daring the
sewers of 1984 - 1987. Preferences among W spp. for bore
diameters, stations, bore opening orientations, and heidnts were
investigated. The influence of bore diameters and bore depth on cell
architecture, the effects of parasitoid lactivity, and aphid prey

selection were also investigated.

pre-e)
of pi
mater
stuo‘y

Varlc

trap
late
cutt
neat

i‘ECt

are
of
DES

gr!

till

CU

io

3t

METHODS AND MATERIALS

Many solitary bees and wasps construct brood cells in
pre-existing natural cavities such as beetle borings or in excavations
of pithy stems and twigs of Sambugus and Jng1ans. Artificial nesting
materials are also acceptable and provide a convenient approach to
study nest architecture, nesting activity, provisions and parasites.
Various materials have been used as artificial nests and included
bamboo, glass tubes, plastic straws, cuttings of twigs and stems, and
trap nests. Condensation in glass tubes and plastic straws make these
materials ineffective. Bamboo has a varying bore diameter and
cuttings of twigs and stems are split with great difficulty. Trap
nests used previously by various researchers consisted of snail
rectangular pieces of wood with holes drilled into their longitudinal
axes. The longitudinal holes (bores) varied in depth and diameter and
are analogous to natural cavities used as nesting sites. Trap nests
of clear straight-grain pine were split in half lengthwise, exposing
nest contents with relative ease, especially if bore diameters were
greater than 3.2 mm.

Trap nest construction techniques for this study were similar to
those described by Fye (1965a) and Krombein (1967). Pine boards were
cut into trap nests (19 x 19 x 140 mm) with holes drilled
longitudinally to depths of 60 and 120 um. Bore diameters and bore
depths varied seasonally, dependent upon prior experience and current

study focus.

it
with 11
height:
ground
increml
of i t:

in all

nests

Split:
selcm
were 3
”Win

blocxs

Bore diameters used in these studies ranged from 1.6 to 9.6 run
with increments of 0.4 or 0.8 mm. Trap nests were usually placed at
heights convienient for making field observations. Heights above
ground level usually ranged from 50 cm to 200 cm with 25 cm
increments. In one experiment. trap nests were distributed at heights
of 1 to 9 m with one meter intervals. Bore depths of 120 mm were used
in all studies with the exception of investigations carried out in
1986 when 60 and 120 mm bore depths were used with equal frequency.

Several problems encountered in early studies were resolved with
modifications of trap nest construction techniques. Snail bore trap
'nests (1.6 um - 3.2 m) were split with some difficulty. The
splitting plane frequently did not intercept the bore. since it was
seldom parallel to the long axis of the trap nest. These problems
were solved by the use of mm trap nests. Several steps were
required for their construction. A band saw was used to cut trap nest
blocks length-wise into two sections with dimensions nominally 6.4 x
19 x 140 mm and 12.6 x 19 x 140 mm. A drill-guide channel was routed
in a longitudinal face of the larger section. Trap nest sections were
bound together with masking tape and drilled to appropriate depths and
diameters with high speed twist-steel bits.

Pre-split trap nest sections did not fit well together.
Irregularities across split surfaces admitted light and excess
moisture, both detrimental to trap nest use. These difficulties were
eliminated by modifying a technique from Krombein (1967). Pre-split

trap nests were coated with melted paraffin and then re-drilled to

their

paraff
suriac
lllust
into :
nests.
secure
bundle
staple
study

in ii

their appropriate bore diameter and depths. Re-drilling removed
paraffin that blocked the bore and produced exceptionally smooth bore
surfaces. Steps in construction of pre-split trap nests are
illustrated in Figure 1. Completed trap nests were bound together
into bund1es of nine (3 x 3), twelve (3 x 4), or twenty (4 x 5) trap
nests. Cotton cord, rubber bands and plastic strapping were used to
secure individual trap nests in bundles. Fye’s (1965a) design for a
bundle carrier was used to place bundles in the field and fence
staples were used to attach bundles to the trunks of trees in the
study area. A typical trap nest bundle is illustrated in Figure 2.

An individual tree with attached trap nest bundles is called a

antinu-

 

 

 

 

 

 

 

 

 

.i

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

D. Pro-split trap neat with bore drilled out.

‘3

c

15

g

D

O

O.“

3; El

__£!
.0

as;

00
=

fiii:

::II

B

as

32..

.u

‘=15

.54!

“—0

'3')

8

La

Q...

‘0

0

i;

h

E?

I:

0

3!

n rap nest face:

. A 6.4 an
t d i t

Figure 1. Construction of pre-spllt trap ne
B

11

 

Figure 2. A 3 x 3 bundle of trap nests ready for distribution into
the field.

I)!

'11
(I)

12

Trap nest stations were established in a maxed hardwood forest
edge between a small red pine plantation and an old field. The long
axis of the edge runs from north-west to south-east. Bundles of trap
nests were positioned so that bore openings faced north-west,
north-east, south-east, and south-west. Trap nests were arranged in
bundles (Figure 3) with bore openings of adJacent trap nests facing
opposite directions, presenting either regular or randomized patterns

of drilled and blank trap nest faces.

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

0 all 9
1. O I
to e o
o . . O
. 0' '
l- - 9' ‘
.0. 9
e 9' '

 

 

 

 

 

 

 

 

 

 

 

 

Figure 3. Distribution of bore openings in 3 x 3 and 4 x 5
trap nest bundles illustrating opposite ends of
representative bundles.

cor
nee
A i

of

an:
09:

COT.-

stu

anc
fen
vit

(b)

the
tra
and

Pro

13

The distribution of bore sizes in trap nest bundles varied
considerably from year to year. Bundles for 1984 contained nine trap
nest of a single bore diameter. Diameters were 3.2, 4.0, and 4.8 m.
A 1985 bundle consisted of twenty trap nests with two trap nests each
of bore diameters 1.6 - 8.8 nm (mum increments). Bundles for the
1986 season were 3 x 4 units of three bore diamter classes (1.6, 2.0,
and 2.4mm; 2.8, 3.2, and 3.6 mm, and 4.0, 4.4 and 4.8 mm) and two bore
depth classes (60 and 120m). Bundles for 1987 were 3 x 4 units
consisting of trap nests with bore diameters of 2.4 to 7.2 m with 0.8
ml increments. The number of trap nests set out for trap-nesting
studies from 1984 throudi 1987 was 4811.

Several techniques were used to rear achit material: (1) trap
nests were placed indivimally into 200 mi polycarbonate rearing tubes
and stored in an unheated garage over winter. (2) pre-pupae were
removed from their cells and transfered to 2 or 4 dram vials lined
with lens paper and (a) stored over winter in an unheated garage or
(b) stored in a refrigerator at 3.5 degrees C for 60 to 90 days.

To gather data on aphid provisions, trap nests were removed fran
the field within one to two days of closure and were replaced with
trap nests of similar bore diameter. Closed trap nests were opened
and data taken on number of cells, cell types, cell lengths, and cell
provisions. The contents of each cell were removed, aphids counted
and the food stores with the wasp egg or larva were transferred to a
rearing vial. Two aphids from each provisioned cell were removed for
identification. Nam prepupae were placed in a cold box at 3.5
degrees Centigrade for 90 days. These materials were removed to

anient lab conditions for rearing of adilts.

All!

The

fat

IQ

iii

pie

line

re;

PI!

COT

Col

thl

Ciel

tile

on

Var

Pin

Pia

act

(1’

14

The study area is a mixed forest edge between a pine plantation
and an old pasture on the most northern portion of Concordia’s campus.
The area is bordered on the east by highway U.S. 23 and on the west by
faculty housing. The plant cover of the study area is given in Table
1. An intermittent stream lies within the edge and an old fence row
roudnly parallels the northern side of the stream. The edge faces
south-southwest and receives maximum sunlidnt in mid-afternoon. Trees
in the area have reached maturity and some are senescent. The
plantation consists of red pine with white pine scattered at the edge.
The edge consists primarily of walnut and ash, with other hardwoods
represented occasionally. A variety of shrubs and herbs are also
present.

Malaise trap collections during the sumners of 1982-1983
confirmed the presence of W on the canmus of Concordia
College, Ann Arbor, Michigan. Preliminary studies were conmcted at
this time to identify an area for intensive W studies and to
develop acceptable trap-nesting material. Pine trap nests with bore
diameters of 3.2, 4.8, 6.4, 8.0, and 9.6 m were placed in three areas
airing the sumner of 1982, but no mm: were reared out. The
variety of nesting materials was expanded for summer 1983 and included
pine trap nests, W stems, cuttings of Rhys, and m; and
plastic straws. Five areas were surveyed and presumed W
activity was noted in each one. Three areas showed minimal activity

(1, 1, and 4 W trap nests respectively): a fourth had 12

trap ne:
for nor:
“in
lboervai
For the
iollovll
opened
ilpeS.
wheneve
Closlre
F00c! st
Vials
identii

a Vaspl

15

W: trap nests and a fifth (area 11) prodiced 27 W

trap nests. Due to these results area II was selected as the location
for more intensive studies of W trap-nesting biology.

Trap nests were placed in the field in late May and early June.
tbservations of W activity were made throughout the manner.
For the 1984-1986 seasons trap nests were removed from the field
following the cessation of provisioning activities. Trap nests were
opened and data were gathered on nest architecture including cell
types, nunber of cells and cell dimensions. Materials were reared
whenever possible. For the 1987 season trap nests were removed upon
closure and data were taken on nest architecture and aphid provisions.
Food stores and wasp eggs or larvae were transferred to snail glass
vials and adults were reared the following spring. Species
identifications were based upon reared materials or unique features of

a wasp’s trap-nesting biology.

16
Table 1.

 

Plant cover of trap-nesting study area.

 

EinusstnonisL- WWI.
Einus.sxixestris.L DION: cubna Muhi

Shrubs
mapp- Rinasp
WWI. Rosaspp
Loninra Blames»
mun Illuminati
mamnus maximum
Rhusradirmnnio lines»

NM“)
Map.
QalJmmnineL- Medals!»
81mins SanminarLacanadensiaL
Simmer.
Innuendo. Solannmsp.
WWI. Solidamspp
Miss» minimal»
Tammi.
mninmlefoiiam) WWI-

 

nun
iii
ne:
01

till

FYI

1r:

tw

01

RESULTS AND DISCUSSIUI

Bore Diameter Preferences Among Sympatric W mp.

Fye (1965b), Kronbein (1967), and Vincent (1978) reported on a
mmber of W spp. and gave data on bore diameters selected.
Vincent (1978) noted 2. W (Say) reared from a 1.5 an bore trap
nest and 2. W Vincent from two 1.5 nm bore trap nests. All
of these authors reported trap nest bores used by 2. sum and
their pooled data are marized as follows: 20 trap nests - 3.2 mm
bore: 83 trap nests - 4.0 In bore: and 3 trap nests - 6.4 ml bore.
Fye (1965b) reported that 2. “111mm preferred 6.4 III bores and
Krofiein (1967) noted 2. W frm four 3.2 m borings, and
two 4.8 II borings.

It is questionable whether these reports are truly indicative of
bore diameter preferences of W map. or represent artifacts
die to the bore diameters selected by the investigators. Bore
diameters most cosmonly made available by these authors were 4.8 u or
greater, with increments of 1.6 an. Fye used 6.4 and 8.0 In
drillings: Irwin also used these sizes and included a few 3.2 nm
bores. Ratios or actual frequencies were not reported. Vincent is
the only author to report use of bores as snail as 1.5 I.
W are snail wasps (4 - 9 mn long) and bore diameters used in
general trap-nesting survey studies may be inappropriate for studies

17

18

focused on this genus, because bore sizes have been too large to be
used effectively by the maJority of these snail wasps.

Bore diameter preferences among W spp. were
investigated from 1984 throudi 1987. In 1984, 432 trap nests with
bore diameters of 3.2, 4.0, and 4.8 up were available. Diamters were
represented with equal frequencies - 144 trap nests of each bore
diameter. Trap nests were bundled into 3 x 3 units and distributed at
16 stations, 3 bundles per station. The following frequencies of trap
nest use by W mm: were confirmed with reared
materials: 3.2 an bore - 19 nests: 4.0 In bore - 9 nests: 4.8 In bore
- 13 nests. An additional 153 trap nests were distributed at a
W site about 100 yards distant frsn the primary study area.
Bore diameters were 3.2, 4.0, 4.8, 6.4, and 8.0 am. W
W was confirmed in 13 of these trap nests: 3.2 in bore - 4
nests, 4.0 In bore - 3 nests, 4.8 m bore - 5 nests, and 6.4 an bore -
1 nest.

Since all available bore sizes were used in the 1984 study, the
nunber of bore sizes for 1985 was increased and ranged frsn 1.6 to 8.8
nm with increments of 0.8 nm. One hundred twenty-eight trap nests of
each bore size were used. Two trap nests from each of these 10 bore
classes were bound together prodicing bundles of 20 trap nests each.
Sixty-four bundles were distributed, 4 at each of 16 stations.

W spp. were confirmed in trap nests with bore diameters
ranging frcml 1.6 to 4.8 m in the following frequencies: 2. mm
(Say), 1.6 llll bore - 2, 2.4 nun bore - 3; E. m Snith, 3.2 an

19

bore - 2, 4.0 nm bore - 2, 4.8 m bore - 3: 2. mm Dahlbom,
1.6 iilll bore - 1, 2.4 m bore - 5: and 2. 5111mm Dahlbom, 2.4 In
bore - 1.

These data suggested that W spp. partition nesting
sites on the basis of bore diameter and that preferred bore diameters
were less than 6.4 In. To increase the frequency of acceptable
nesting material and bore categories, the maxinmm bore diameter used
in 1986 was 4.8 III. Nine bore diameters were used, ranging from 1.6
mm to 4.8 mm with 0.4 mm increments. One hundred twenty-eight coated,
pre-split trap nests of each bore diameter were prepared and assembled
into 96 3 x 3 bundles. Bundles were of 3 classes: class I, 1.6 - 2.4
mm bores: class II, 2.8 - 3.6 mm bores: and class III, 4.0 - 4.8 mm
bores. Each bore dim-eter was represented 3 times in their respective
bundles. Bundles were distributed to 24 stations, 4 per station.

One hundred forty-seven trap nests were provisioned by
W spp. and species confirmations were made for 104 trap
nests by reared materials or trap nest biology. Four species of
M were reared: E. W (Say), 6 trap nests - 8 males
and 3 females: 2. W Vincent, 47 trap nests - 17 males and 42
females: 2. gnsp1da1us Smith, 33 trap nests - 58 males and 13 females:
and 2. mm Dahlbom, 18 trap nests - 17 males and 22 females.
Forty-five trap nests, presumed to contain Eassa1gegus, suffered cell
failure or rearing losses and no adults were reared. These trap nests

were initially classified as provisioned by unknown W spp.

110

US

(11

tr

(11

th

to

<<

20

However, subsequent examination of trap nest closure materials were
used to identify thirteen unknown trap nests as being provisioned by
E. W Vincent.

Table 2 smarizes data for 1984-1986 on frequency of bore
diameter availabilty and selection as nesting sites by five
W spp. The Kruskal-Nallis test for differences in ranks of
trap nest bore selection by four of these species (B. W, 2.
W, E. W, and 2. W13) is very significant (ll
8 120.9749, df = 3, p «.0005). The chi-square (I) test for
differences of bore diameter selection by 2. W (based upon
three bore diameter classes: 2.0 - 2.8, 3.2 - 4.0, and 4.4 - 6.4 m)
is very significant (x2 - 15.2583, a: = 2, p <.ooos> The t(Ii) test
for differences of bore diameter preferences between 2. 5mm.“ and
E. W is also very significant (t ‘= 7.4316, df = 116, p
«.0005).

21

Table 2.

 

Bore diameter selections for five We spp., 1984-19$.

 

Bore dimters (I)

 

1.6 2.0 2.4 2.8 3.2 3.6 4.0 4.4 4.8 5.6 6.4
Bore diameter freqiencies

 

256128256134$128491%4m128158

Species

mm 3 1 7 0 0 0 0 0 0 0 0
mm“: 18 20 9 0 0 0 0 0 0 0 0
m 0 1 4 6 32 7 19 3 21 0 1
“1111mm 2 3 13 2 2 2 0 0 0 0 0
31m 0 0 i 0 0 0 0 0 0 0 0

 

Pa

81

22

In 1987 the focus of these studies was narrowed to E. W.
Paraffin coated, pre-split trap nests of seven bore diameters (2.4 -
7.2 nm, with 0.8 lllll increments) were used. The frequencies of bore
sizes made available as potential nesting sites were as follows: 2.4
in - 211: 3.2 lilli - 307: 4.0 nm - 307; 4.8 m - 307: 5.6 nm - 211; 6.4
in - 211 and 7.2 - 211. 2. mm used bores in the following
frequencies: 2.4 m - 9: 3.2 m - 37: 4.0 nm - 28: 4.8 um - 8: 5.6 [III
-1, and 6.4 m - 1.

2. My: trap nest selection data from 1984—1987 were pooled
for analysis and are given in Table 3. Due to insufficient data in a
few bore diameter classes, bore diameter selection data were
recsmined into five classes. The expected frequencies of trap nest
selection were based upon relative frequencies of trap nests in each
bore class and the expectancy that the ratio of trap nest use would be
equal in all classes. The expected frequencies of trap nest use in
respective bore classes were: 27.17, 39.50, 39.50, 39.50, and 32.33.
The observed frequencies of selection in these respective bore
diameter classes were 14, 75, 54, 32, and 3. Expected and obarved
frequencies of bore selection are significantly different (Fimre 4,
x2 - 71.6456, a: =- 4, p «.0005) indicating that 2. W prefers

trap nests with bore diameters from 2.0 - 4.8 lllll.

23

Table 3.

 

2. mm: bore diameter selections, 1984-1987.

 

 

 

 

Bore diameter Dineter Selection

class (I) freqiency fremencv
2.0 - 2.4 595 14
2.8 - 3.2 $5 75
3.6 - 4.0 $5 54
4.4 - 4.8 $5 32
5.6 - 6.4 708 3

 

CODmCDNfi-rl

24

Passaioecus cuspidatus
Bore Selection Frequencies, 1984-i887

 

0
2.0-2.4 2.8-3.2 3.6-4.0 4.4-4.8 5.6-8.4

Figure 4.

Bare Diameter Classes (min)
.Expected Frequency fiflbserved Frequency

Expected and observed frequencies of bore diameter
selection DY 2am: W. 19344937-

 

 

25

A possible factor influencing bore diameter selections among
W spp. is wasp size. A relatively simle index to wasp size
is head width. iiead width measurements, to the nearest 0.1 an, were
taken frsn samples of ten females of E. W, B. W,
and E. W. The respective mean head widths for these samles
were 1.46, 1.19, and 1.0 cm. W spp. partition nesting sites
on the basis of bore diameter and the size of the wasp may limit the
minimum acceptable bore diameter. An additional factor in this regard
may be the size of aphids selected as prey. Aphids are usually
carried in the mandibles with the prey’s body lying below the wasp’s
head. Under these circumstances the dorsal-ventral dimension of the
wasp’s head plus an aphid will be greater than head width and will

certainly influence acceptable bore diameters.

Trap Nest Meidnt Selection in W mp.

Trap nests were distributed at several heidnts to determine those
acceptable to W spp. and to determine if height was a
significant factor in the partitioning of nesting sites. Trap nest
bundles were distributed at heidnts of 0.5 to 9.0 meters. Intervals
below 2.0 m were 0.25 in. One meter intervals were used at heidnts
above 2 m. Ram mp. used trap nests at all available heidnts.
Pooled data for 1984-1987 are given in Table 4. The frequency of trap
nests used by four W mp. at various heidnts is given along
with the number of trap nests of acceptable bore diameters distributed
at those heidnts.

E. W and 2. 01901119912111: were restricted to heidnts below
3 meters. 2. medians and 2. W used trap nests from a wide

range of heidnts, 1 - 9 m and 0.5 - 7 m respectively. However, these
data do not indicate upper limits of heidnts that midnt be used by
these two mecies. The data are particularly interesting in the case
of 2. am and B. W, the first preferring heidnts above
three meters and the second heidnts below three meters. Chi-smaare
(I) was used to test for differences in heidnt distribution of trap
nests selected by z. W and E. W. Data were pooled
into three height classes for each species and expected frequencies of
trap nest selection were based upon frequencies of acceptable bore
diameters. For 2. mm: heidnt classes were 0.5 - 1.0, 1.25 -
1.75, and 2.00 - 9.00 m: observed frequencies of trap nest selection
26

27

for these height classes were 63, 84. and 17: and respective expected
frequencies of trap nest selection were 71.70, 63.23 and 29.06. The
chi-square (I) statistic for these data is significant (X2 = 12.8819,
df = 2, p < .005). For E. 55:9131u3 height classes were 0.5 - 3, 4 —
6, and 7 - 9 meters: observed frequencies of trap nest selection were
22, 12, and 15: and respective expected frequencies of trap nest
selection were 43.49. 2.75, and 2.75. The chi-square (1) statistic
for these data is very significant (X2 = 96.1107, df = 2, p <<.0005).
These results have implications for future trap-nesting studies.
supporting the notion that height should be considered in distribution
of nesting materials, and suggest that heidnt may be a factor in

partitioning nesting material.

28

Table 4.

 

Trap nest heidnts and frequency of use by four 2mm: mp.,
1984-1987.

 

 

Total 8 of trap nests used (4 offered)! to W mp.

Meidnts trig

(m) nests annu l atus areoi atus cum 1 datus moni l learn is
9.00 48 0 (48) 15 (48) 0 (48)
8.00 48 0 (48) 0 (48)
7.00 48 3 (48)

6.00 48 0 (48) 12 (48) 0 (48)
5.00 48 0 (48) 0 (48)
4.00 48 2 (48)

3.00 48 0 (48) 13 (48) 0 (48)
2.00 764 1 (160) 5 (160) 12 (604) 4 (372)
1.75 586 0 (70) 34 (516) 0 (204)
1.50 860 4 (160) 1 (160) 9 (700) 5 (372)
1.25 586 41 (516) 0 (204)
1.00 908 4 (160) 3 (160) 17 (748) 9 (372)
0.75 586 25 (516) 0 (204)
0.50 860 2 (160) 0 (160) 21 (700) 5 (372)

 

 

 

 

in MFG in parentheses indicate the nunber of trap nests at
heidnts that are of acceptable bore diameter for a mecies.

given

Parasites of W mp.

Parasites of Ea:sa1gegus,gusp1da§us have been reported by Krombein
(1967) and Vincent (1978) and included the chrysidids mam: ms
(Fabricius), mm W (Provancher), and ichneumons M
w m (Cresson). and mm mm (Cremon). Of
nine hundred twenty-eight Passa1g§gus spp. cells provisioned. 88 were
parasitized and 70 parasites were reared. Parasites included mam:
gang“: (Fabricius), Q. 1:1gesggns (Norton), Ir1ghrys1siggz1ag Gribodo.
and m 1111.121: (Cresson) and 2. mm (Cremen).

Table 5 smarizes data on ms megs parasitisn of E.
W for 1984. Of 25 trap-nesting stations, 21 were used by P.
gnsn1gagns and 14 had trap nests parasitized by Q. agngns. Q. agngus
parasitized 35.19% (19 of 54) of all 2. mm trap nests and
17.06% (29 of 170) of all 2. W cells. Stations with hidn
frequencies of W activity (3 or more trap nests per station)
snowed a wide range of parasite activity. Station 3 prodiced four
Paaaa1gggns, nests with 21 cells and no cells were parasitized.
Station 11 prodiced 3 nests and 6 cells, with 10096 nest parasitim and
66.668 of the cells parasitized. Station 8A generated 5 2, gusn1datus
nests *with 26 cells. Three of these nests and 9 cells were

parasitized.

Table 5.

 

Distribution of m: am (Fabricius) among
Eassa1gggus,gusp1datus trap nest stations. 1984.

 

 

 

 

 

Parasitized

Station trap nests cells nests cells
1 4 19 1 2
2 2 5 0 0
3 4 21 0 0
4 0 0 0 0
5 1 2 0 0
6 2 4 1 1
7 4 9 2 2
8 3 5 1 1
9 3 9 1 2
10 3 10 1 1
11 3 6 3 4
12 4 13 1 1
13 2 5 1 1
14 1 2 1 1
15 4 14 1 1
16 1 2 0 0
1A 2 3 1 1
2A 0 0 0 0
3A 0 0 0 0
4A 2 10 0 0
5A 1 1 0 0
6A 2 3 1 2
7A 1 1 0 0
8A 5 26 3 9
9A 0 0 0 0

 

31

Data from 1984 were also examined for bore diameter preferences
of 0. may: and are summarized in Table 6. These data show no
significant difference in frequency of parasitisn based upon bore

diameter differences.

Table 6.

 

Parasitisn of W W trap nests and cells
by mm: mm (Fabricius) by Bore Diameter. 1984.

 

Bore diameter (um)

 

3.2 4.0 4.8 6.4
gu;2?§atys 23 12 18 1
Eéap nes s
Number of
. 78 29 52 10
cel s
8 tra nests 26.09 33.33 27.78 100.0
paras tized
8 cells parasitized 12.82 17.24 44.44 20.0

 

In 1985. two of 25 W nests prodiced parasites: one

mm W from a trap nest of 2. mils and two mm
m Gribodo from a single 2. gym trap nest.

Table 7 gives data on the distribution of parasitisn among
W trap nests for the 1986 season. Twenty-four of 28
stations prodiced W nests and 11 stations had parasitized
nests. m: m was reared from two 2. 3mm: nests, four 2.
mm: nests, two 2. W nests, and one nest of B.
“11.1mm:- 2mm alum (Cresson) was reared from six 2.
mm: nests. six 2. m nests. and one nest of 2.
alumna.

cells

cells

magnesia Poemnia

32

Table 7.
cells

nests

Distribution of mains means and Romania among Passaims

trap nests, 1986.

 

 

Station

01100000.20010020.0.0100272. _

 

— 07010000.01000010.0.0050240. _

_ nxsviasaummqauimmus“ssmnmmmu_

1 an. 1

123assvaemummuemnmwmmnmusxmm

375

Totals 140

 

33

Data frsn 1986 were also examined for the height distribution of
nests parasitized by My: new: and mu 31:11.22:- Q. m
parasitized nests at heigints of 1 to 7 in, while 2. 11m parasitized
nests at heidnts of 0.5 to 9 m.

The distribution of W parasites according to station
for 1987 may be noted in Table 8. Twenty-two of 49 stations were used
as nesting sites by 2. We. Nests at six stations were
parasitized by Mill: m and one station prodiced mm
mm. Of special note are stations 36. and 47 throudn 49.
Station 36, established on May 30. prodiced seventeen E. W
trap nests and sixty-five provisioned cells. As of June 15 four nests
were parasitized. On July 1 three additional stations (47 - 49) were
established nearby, each within 5 meters of station 36. These
stations generated an additional 17 E. W nests, and 59
provisioned cells, but none were parasitized. Trap nests at all of
these stations were distributed at the same heidnts and with similar
orientations. At station 36. 2. W used eidnt trap nests at a
heidnt of 1.75 in. another eidnt at 1.25 m, and one at .75 11:. Among
stations 47 - 49 2. W used seven trap nests at a heidnt of

1.75 m and ten trap nests at 1.25 m.

it i:
distributl
or to sea
shoe tha'
12 and a
prey on
behavior
first i

activit

34

It is questionable whether the unevenness of 0, agneus
distribution at Eassa1gggu§ stations is due to host switching behavior
or to seasonal changes in host availability. Data from 1984 studies
show that 0. gene“: parasitized 2, gnsp1gagus nests as early as June
12 and as late as July 24. These data suggest that Q. annoys could
prey on E. gusn1gagus as long as it is active and that the searching
behavior of n. m is limited to an area where host activity is
first observed. However, other site factors may influence the

activity of this parasitoid.

 

Table 8.

 

Distribution of Parasitisn among W W
trap nests. 1987.

 

Station trap nests cells parasitoids

 

 

p-m

mmmntmflflfl
immmmmwn

H

A
AmfimflbwflwmpONw

immmmmwn

immmmmmn
immmmmwa

manna“:
manna”:

8

b

‘1

p-m
OOOOOOflOO‘IOO-‘QOOOONOOOOHN

(A)
”
QbOF‘HN‘JF‘fl-bNmflfle-‘F‘ONNF‘QF‘UI

488m~m8n$

 

Provisioned Trap Nest Architecture

A general discussion of provisioned trap nest architecture, as
illustrated in Figure 5, follows. The innermost portion of a trap nest
is sometimes left empty and walled off by a W 91m. This
portion of a nest is called the M1 5111. Wed 9:11: consist
of a food-stores mace and partition materials. The length of the
foodbstores space and thickness of the partition comprise the length

of the cell. Infrequently, empty (W) cells are found

between provisioned cells. The outermost portion of the nest is
usually enpty and closed by a partition at the nest opening. This
final cell is the 11331131131: 911.1. and its partition is the 11mm.

Provisioned Cells

Basal Cell 1 \<:\\\k Vestibular Cell
Preliminary Plug \ //—— Closure

Resin Partitions

 

 

 

     
 

 

Figure 5. Nest architecture illustrating basal cell, provisioned
cells, vestibular cell, preliminary plug. resin partitions
and closure.

36

 

Pit

1M

(0

40C

2.12

NC

1115

Par.

Etc

len

Factors Influencing Cell Length and Volume

The raw data for provisioned cell lengths for these three
2am mp. were quite varied and skewed. This should be
expected since one tail of the possible distribution of provisioned
cell lengths is closed (no cell can have a length shorter than 0.00
an) and the open end of the distribution is limited by the actual
length of the trap nest bore (60 or 120 m). 2. mm: provisioned
trap nests of three bore diameters - 1.6, 2.0. and 2.4 run. Data for
provisioned cells frsn these three bore diameters are remectiveiy:
number of cells (49, 73, and 30): range of cell lengths (9 to 70, 7.5
to 41, and 7 to 36 nun): median cell lengths (16.25, 13.19, and 13.19):
and mean cell lengths (19.79 1 12.79, 14.33 1 5.31, and 15.57 1 7.27).
B. W provisioned trap nests in 10 bore diameters ranging from
2.0 to 6.4 nmn. Four bore diameter classes (2.4, 3.2. 4.0, and 4.8 m)
prodiced significant numbers of cells. Data for these four bore
diameters are remectiveiy: nunbers of cells (40, 210, 1m, and 95):
range of cell lengths (8 to 39, 6 to 82, 5 to 101, and 5 to 116 an):
median cell lengths (12.96, 12.73, 10.03, and 10.23 nun): and mean cell
lengths (14.34 + 5.29, 14.17 1 8.30, 15.09 1 17.00, and 20.58 1

26.33).

37

38

beamination of ranges and medians for these respective species
and bore diameter classes show median cell lengths much shorter than
cell lengths at mid-points of remective ranges. in normal
distributions we would expect equal proportions of measurements above
and below the mean. However, the percentage of cell lengths shorter
than remective mean cell lengths for 2. 311911151: were 71.4, 68.5,
and 73.32. For E. gusp1ga1ua these values were respectively 72.5, 61,
80, and 81.5%.

In each of the above cases mean cell lengths are longer than
median cell lengths and variances are exceptionally high. These data
reflect the statistical effects of a small number of provisioned cells
of extraordinary length. These cells were usually found to be. but
not limited to, the last provisioned cell in a trap nest. Frequently
such a nest did not have a vestibular cell, although in a few cases an
extraordinarily long cell was followed by a vestibular cell. To
eliminate the statistical effects of cells of extraordinary length. I
arbitrarily applied a 10’: exclusion role in the analysis of cell
length data. For each species, 10* of the pooled cell length values
from the open end of the distributions were excluded in subsequent

analysis.

[‘6

i'lUl

to

 

39

Based upon the 10% exlusion rule, 2, areg1agus cell length data.
respectively for 1.6. 2.0, and 2.4 mm bore diameter trap nests. are:
number of cells (41, 71, and 25): range of cell lengths (8 to 25.5, 7
to 25, and 7 to 21 mm): median cell lengths (14.17, 13.06, and 12.81
In): and mean cell lengths (15.13 1 3.81, 13.51 1 3.79, 12.68 1 3.26).
Similarly, data for E. W provisioned cells from bore
diameters 2.4, 3.2, 4.0 , and 4.8 m are remectiveiy: number of cells
(38, 195, 164. and 79): range of cell lengths (8 to 22, 6 to 23, 5 to
23. and 5 to 23): median cell lengths (12.89. 11.93, 9.90. and 9.825
Inn), and mean cell lengths (13.43 1 3.19, 12.39 1 3.45, 10.33 1 2.38.
and 9.47 1 2.72 mm).

One way analysis of variance for differences in cell lengths
associated with differences in bore diameter were significant for 2.
argg1a1us (F 8 3.30337, df = 134, p (.01) and for 2. gugp1ga1us (F -
19.51697. df = 472, p (.001). In the case of E. W, data
were too scanty for the application of normal based statistics, but
the Kruskal-Nallis rank order test for differences in cell length data
was significant (ll - 25.7153, df = 4, p (.0005). These results
indicate that for each of these W mp. an inverse
relationship exists between bore diameter and cell length. A surmnary

of these analyses are given in Table 9.

40

Table 9.

 

Mean provisioned cell lengths for three W mp. frem pooled
mics, 1984-i907.

 

W mp. provisioned cell lengths“ (n)

Bore
diaeter
(I) areolatus (ll) cumidatus (ll) monilioornis (ll)

 

 

 

 

1.6 15.13 1 3.81 (41) -- --
2.0 13.51 1 3.79 (71) - 14.17 1 2.72 (3)
2.4 12.68 1 3.26 (25) 13.43 1 3.19 (38) 12.47 1 2.95 (38)

2.8 - - 9.81 1 2.09 (8)
3.2 -- 12.50 1 3.59 (195) 9.93 1 1.27 (7)
3.6 -- - 7.42 1 .61 (6)
4.0 -- 10.47 1 3.11 (164) --
4.8 - 10.07 1 3.37 (79) --

 

I Ten percent of pooled sqie values frem stewed end of
distribution were excluded in analysis.

With the application of the 10’: exclusion rule mean values are
clearly more representative cell length measurements and demonstrate
an inverse reiationdnip between bore diameter and cell lengths.
However, they do not reflect the extreme variation observed in
provisioned cell length and we can only meculate on possible causes
for such variation. For examle, one possible cause could be a
declining prey population. As the prey nnmers decrease, additional
time and energy are expended daring provisioning. In this case an

optimal closure or partition construction strategy would be the

red

(0

tie

len

for
inc
ler

]

DU D... 00 \

41

rediction of the time required to move frsn the trap nest bore opening
to the partition or clomre. A partition or closure establisined
closer to the bore opening will prodice a cell of dimroportionate
length and volume.

Results of regression analysis of bore diameter and cell length
for three W: mecies are given in Table 10.; These analyses
indicate that larger bore diameters result in shorter provisioned cell

i engths .

Table 10.

 

Simle linear regression of bore diameters and mean cell lengths for
three We app.

 

Co-efficient of

 

 

 

 

Species Regression l ine correlation Significance
2. W Y - -3.06)( + 19.90 -.98 P a .10
20 W Y 8 -1051x + 17.07 -997 P < .05

E. W Y - -4.01X + 21.99 -.97 P < .01

 

 

 

905

It

42

Banks (1971c) mggested that the availability of nesting sites
was one of the factors limiting populations of aculeate Mymenoptera.
If this is true, it should be possible to identify strategies used to
optimize available nesting materials. One of these strategies could
be the decrease of cell length as bore diameter increases. Fye
(1965b), Kronbein (1967) and Vincent (1978) gave bore diameter and
cell length data for E. W. Kraflnein (1967) reported that
fifty-eidnt provisioned cells from 3.2 min bore diameter trap nests had
a mean length of 16.3 nm and a range of 8 - 52 all. Four cells frem
4.8 on bore diameter were 7, 8. 13. and 126 in long and one 6.4 an
bore diameter trap nest had four cells 6, 7, 7, and 9 on long
remectively. Fye (1965b) reported a 6.4 an bore diameter with four
cells, with a mean cell length of 15 Ill. Vincent (1978) reported data
from 83 soda straw nests with 4.0 nm bore diameters. One
hundredceleven female cells had a mean length of 10.09 1 2.19 m and
one hundred-ten male cells had a mean length of 8.82 1 2.16 illll.

Data frem [rowein (1965b) and Vincent (1978) suggest that
W mp. would optimize nesting material by decreasing cell
length as bore diameter increases. To test whether W mp.
would optimize their use of bore volume. trap nests of several bore
diameters were made available as nesting sites. Provisioned cell
length data were collected frcmn trap nests provisioned by four
We spp.: annuiatus. annulus. W. and Minimum.
Adequate samle sizes were obtained by pooling data frsn 1984-1987 for
E. W and E. mm. Data for E. mm were
sketchy.

hum

43

Previous research on We makes no reference to cell
volume, and no anaylses have been done on relationships between nunber
of prey per cell, cell length, cell volume and bore diameter. If
W spp. made maximum use of available bore mace we could
expect constant volumes for cells from different bore dimneters. This
hypothesis was evaluated by determining cell volumes from mean cell
lengths and bore diameters for P. W, 2. gym, and 2.
W. The results of these calculations are given in Table
11, and slnow that mean cell voltmne increased along with increasing

bore diameter for each species considered.

Table 11.

 

Bore diamters and provisioned cell volumes for three W mp.

 

W mp. provisioned cell volumes (m3)

 

 

 

 

Bore

(II) areol atus cumidatus moni l icornis
1.6 30.42 1 7.66 -- ~-

2.0 42.44 1 11.90 -- 44.52 1 8.54
2.4 57.36 1 14.77 60.76 1 14.43 56.41 1 13.34
2.8 -- -- 60.41 1 12.87
3.2 -- 100.53 1 28.87 79.86 1 10.18
3.6 -- -- 75.53 1 5.54
4.0 -- 131.57 1 39.08 --

4.8 -- 182.22 1 60.98 --

 

 

 

91‘

ha:

th

59

CC

I!

44

A related question is whether increased cell volume is associated

with larger nunber of provisions. Aphid provisioning data were
analyzed for 3.2 and 4.0 mm bore trap nests provisioned by 2,
W with Mung M (Davis). Data for cells with
extraordinary length and/or cells in which significant larval feeding
had occurred prior to examination of cell contents were excluded frcml
this analysis. Data were available from 58 cells of 3.2 m bore and
59 cells from 4.0 nm bores. Ranges and means for number of aphids per
cell, cell lengths. and cell volumes were determined for these
remective bore diameter classes and are presented in Table 12. The
t(II) test shows no significant difference in the numbers of aphids
provisioned in 3.2 and 4.0 nun bore trap nests. However, differences
in cell lengths and volumes 'were signficant. As bore diameter
increases, cell lengths decrease and cell volume increases. No data
were collected on actual volumes of aphid provisions but a relative
index to utilization of available mace is cell volume (m3)/aphid.
The index for 3.2 on bore trap nests is 2.895 and for 4.0 m bores.
3.336. If an equal mean volume per provisioned aphid is assumed for
these bore diameters, these indices slow a more efficient use of the
3.2 In bore. These results agree with the general subJective
observation that the free space above the aphid provisions was larger

in 4.0 m bores.

45

Table 12.

 

An analysis of number of aphids [51103 monazggg (Davis)] provisioned per
cell, cell lengths and cell volumes in 3.2 mm bore and 4.0 mm bore trap
nests provisioned by E. guspigatgs, 1987.

 

tiII) statistic

 

 

 

 

 

 

 

Bore Range Mean t value df prob

t of aphids 3.2 22 to 66 35.81 1 10.25

per cell 1.3098 115 p >.05
4.0 14 to 74 38.67 1 13.03

Cell 3.2 7 to 23 12.89 1 3.29

length 4.2796 115 p (.0005

(an) 4.0 6 to 23 10.27 i 3.28

Cell 3.2 56.30 to 184.98 103.67 i 26.49

volume 3.9331 115 p (.0005

(.13) 4.0 75.40 to 289.03 129.02 1 41.19

 

V0

46

Scatter plots of numbers of aphids provisioned per cell and cell
volumes for 3.2 and 4.0 III) bore trap nest are given in Figures 6 and
7. Siiiple linear regression of cell volumes on number of aphids
provisoned per cell gave the respective regression equations: (Y =
54.88 + 1.36X, r = .53) for 3.2 mm bores and (Y = 86.75 + 1.09X, r =
.35) for 4.0 lilli bores. The correlation coefficients for these data
indicate what cell volume is weakly related to numbers of aphids
provisioned. In the case of 3.2 mm bores, 25% of the variance of cell
volume is accounted for by the variance of numbers of aphid
provisions; in 4.0 m bores only 12.25% of the variance of cell volume
is accounted for by the variance of the numbers of aphid provisions.
Cell volumes are not closely related to the numbers of aphids

provisioned.

47

.hooa .mumo: nous upon
EE N.m “adamamnmu .m 90w AmEEv moE:_0> __ou Dec
__00 sod Ooco_m_>oLa mU.£Qo «o guess: «0 uo_a souuoom .w sham—m

:3 sum 3533?... 2.2.5 .3 33:52

=~ ca on a. an cu
- _ _ _ ha on

, a new .am
a n a annu a .ea

a a an .3
.2:
.e:
.92

.2:
.3.—

.2:
.2:
a
a .2:

 

 

 

.33 .33: an: 33 E... a.” 33333 .._ 3..
$523 :3 can :3 3.. 353.33.. 322.

(Jlflh—n—n ZDrEI—nillEEdD

 

“HIE-ulc-i Flu?! Che-Ill Ila-aft Ill-ll Ill

 

 

48

.hmou .mumoc moss upon
65 o.v HJMMUAmflmu .m Law nmEEv mmE:_O> ._00 new

__oo boa noco.m_>0sa mU.£Qo no 9025:: «o uo.a gouusom .fi ous®.m

=3 .2 3:333... 322. .3 33:52

3

 

 

.32 2.33: an: 33 SE a... 33333 .m 3..
muss—o: :3 can :3 can. 35339:. 23.3:

3

.8

.2:
$2
.3—
.2:
.8—
53
.eau
.9:
.Su
.Su

 

gs

Gill—u— DD—DEU

49
In addition to bore diameters, several other factors could
contribute to variations in lengths of provisioned cells. Wasp
behavior related to the placement of cell partitions and ciowres,
could be altered by age, prey availability, proximity of prey, prey
switching, weather conditions, competition for nesting sites, and the
activity of parasites. To determine whether extraordinary cell
lengths were a result of wasp senescence, cell length data for trap
nests with known closure dates were examined from 2. My: trap
nests of 1984 and 1987. If extraordinary cell lengths were a result
of senescence, such lengths would be noted most frequently in
late-season nests. Provisioned cells of extraordinary length (24 -
116 mm) were found in trap nests provisioned throughout 2. gnspidatug
nesting season, and in all but three cases the extraordinary cell was
the last cell provisioned (Table 13). Senescence can thus be excluded

as a principal factor contributing to increased cell length.

50

Table 13.

 

Seasonal distribution of Eaaaaigggg§_ggsplgatu§ cells of extra-
ordinary length.

 

Date of Bore Cell length (mm)
closure (mm) 1 2 3 4 5 6 7 8 9
6—18-84 4.0 11 11 94*

6-18-84 4.0 15 101*

6-20-84 4.8 116*

6-21-84 4.8 8 108*

7-09-84 4.0 7 56*

7-16~84 4.8 17 24* i7 56*

7-16-84 4.8 14 14 72*

8-07-84 4.8 14 12 78*

6-15-87 4.0 10 10 72*

6-19-87 4.0 13 97*'

6-19-87 5.6 7 7 101*

7-03-87 3.2 19 19 12 67*

7-06-87 4.0 7 7 8.5 14 7 7 7.5 13 46*
7-08-87 4.0 13 10.5 15.5 10 10 67*
7—08-87 3.2 47*

7-08-87 3.2 16 47*

7-08-87 4.8 10 13 77*

7-08-87 3.2 23 25* 51*

7-10-87 4.0 12 15 14 59*

7—10—87 4.0 10 8 8 94*

7-20-87 4.8 9 105*

7-20-87 4.0 10 9 46*

7-20-87 4.8 35* 88*

7-20-87 4.8 12 12 91*

7-20-87 4.8 13 13 44*

7-24-87 4.0 23 75*

7-27-87 3.2 12 16 17.5 82*

 

*: extra-ordinary cell length

 

of pa
91 id!
1209'
Para
veil

trap

51

Another possible cause for increased cell length is the activity
of parasites. Data from E. cumidatus nests of 1984 were examined for
evidence that 0. am activity contributed to increased cell
lengths. Bidlteen of 53 provisioned trap nests were parsitized.
Parasitized trap nests contained a total of 59 cells and 27 of these
were parasitized by Q. m. Cell length data for these parasitized

trap nests is given in Table 14.

Table 14.

 

Cell length data fra 1984 W W nests parasitized
by 0. m.

 

Date of Bore Cell length (I)
closure (I) 1 2 3 4 5 6 7 8 9 10

 

6-14-84 3.2 31* 10 24
6-17-84 3.2 12* 11 14 12 12 10* 10 11
6-18e84 3.2 18* 20* 27* 24*
6P30-84 3.2 18 19 13 9*
7-16-84 3.2 22* 15
7-24-84 3.2 27 8 12*
6-18-84 4.0 32* 6 9*
6-19‘84 4.0 25* 9
6-21-84 4.0 11 15*
6-26-84 4.0 12 11 9 9 8* 8
7-03-84 4.0 8* 10
6-12-84 4.8 7*
? 4.8 10* 10*
6-21-84 4.8 8* 108
6-30-84 4.8 10* 10
7-16-84 4.8 14* 14 72
7-16P84 4.8 17* 24 17* 56
6-18-84 6.4 6* 6 6* 6 5* 6 6 6 5 21

 

*: parasitized cell

bore

101

cell
94,
lens
date
110;
cel!
11a:

15.

52

Lengths of provisioned cells from all trap nests of respective
bore diameters were as follows: 3.2 lllll (6 to 31 m, n = 79): 4.0 lllll (6
to 101 m, n = 29); 4.8 Ill! (5 to 116 um, 11 = 52); and 6.4 Illll (5 to 20
ill. n = 10). Cell length data for eidlt of 170 cells (approximately
5% of all values) were excluded in the analyses of these data. No
cell lengths were excluded from 3.2 illli bore data, 3 cell lengths (56,
94, and 101 in) were excluded frail 4.0 lllll bore data, and 5 cell
lengths (56, 72, 78, 108, and 116 m) were excluded frail 4.8 lllll bore
data. No parasitized cells were noted among excluded cells. In each
trap nest bore class, mean cell lengths were determined for: all
cells, cells from trap nests free of parasites, cells from parasitized
trap nests, and parasitized cells. These data are sumarlzed in Table

15.

53

Table 15.

 

Cell length data for cells in: non-parasitized trap nests and parasitized
trap nests provisioned by E. 1211211181113. 1984.

 

llean Provisioned Cell Lengths (I), Standard Deviations,
and Idler of Cells in Bach Class

 

 

 

 

 

 

 

Dore All Trap llests Trap llests Free Trap llests with Parasitized
(I) in Bore Class of Parasites Parasites Cells Only

7 13.494 12.421 16.00 18.727
3.2 s 5.207 3.911 6.742 6.916

n 79 57 22 10

1'1 12.192 12.8 11.8 16.167
4.0 s 6.102 4.833 6.747 9.263

n 26 10 16 6

i 9.787 8.861 12.818 11.625
4.8 3 4.110 3.376 4.783 3.638

n 47 36 11 8

31' 7.3 - 7.3 5.667
6.4 s 4.584 - 4.584 .471

n 10 - 1 3

 

51- lean; s a standard deviation; n = niner in class.

54

Thoudi sanple sizes are relatively snail these data show that
parasitiall of 2. 21121111111: cells by Q. new: results in increased
cell lengths in trap nests with bore diameters equal to or less than
4.8 mm. The t(II) test for differences in mean cell lengths between
trap nests without parasites and trap nests with parasites was
significant for 3.2 lllll trap nests (t = 2.343, df = 77, p (.025) and
for 4.8 an trap nests (t = 2.5563, df= 45, p (.005). Parasitized
cells frail 3.2, 4.0, and 4.8 lilli bores had cell lengths which were
respectively 51%, 26*, and 31% longer than cells from non-parasitized
trap nests. In a single 6.4 an bore trap nest parasitism did not

result in increased cell lengths.

Bore Depth and Cell Architecture

it was noted from trap-nesting studies of 1985 that small bore
trap nests (1.6 m), with depths of 120 um, had basal cells of
considerable length (84 to 114 Hill). Additionally. large bore trap
nests (3.2 - 4.8 lllli) occasionally had long vestibular cells. If
availability of nesting sites were a limiting factor in provisioning
activity, a wasp should maximize its use of available space in a trap
nest. However, several factors could mitigate against maximization of
bore volume. The distance traveled from the trap nest bore opening to
the interior of the trap nest could represent a considerable
expenditure of time and energy during cell provisioning and closure.
If an aphid colony were some distance from the nesting site or if
aphids were in short supply, a strategy that descreased the real time
of provisioning activity would be optimal. To study the response of
W spp. to decreased bore depth, trap nests for the 1986
season were provided with bore depths of 60 and 120 an. Data were
collected on lengths of basal cells, provisioned cells, and vestibular

cells: and nullbers of cells provisioned.

55

 

56

Sixty-five W spp. trap nests with bore depths of 60 illll
had basal cells with a mean length of 4.05 lilll. Sixty-elmt
W spp. trap nests with bore depths of 120 illll had basal cells
with a mean length of 22.09 nm. The Z — approximation of the
liann-lihltney U ranks test for these data was very significant (2 =
3.492, p (.0003). Data for basal cell lengths from We spp.
trap nests are suillarlzed in Table l6. A decrease in bore depth was
accolipanled by a decrease in mean basal cell length. W spp.
were clearly sensitive to changes in bore diameter and depth. As bore
diameter decreased, 8 longer portion of the trap nest bore was unused.
Trap nests with bore diameters less than 3.2 m most clearly

demonstrate this phenomenon .

Table 16.

 

iiean basal cell lengths from eiglt bore diameter classes and two bore
depth classes. Pooled data frml W trap nests, 1986.

 

Bore depths (nun)

 

 

 

 

 

Bore Diameter
diameters (um) 60 (ii) 120 (ii) Class means
1.6 12.93 (15) 38.00 (11) 23.54 (26)

2.0 0.00 (17) 23.72 (18) 12.20 (35)

2.4 4.60 (15) 30.94 (16) 18.19 (31)

2.8 0.00 (3) 17.38 (6) 11.89 (9)

3.2 0.00 (6) 7.50 (4) 3.00 (10)

3.6 0.00 (6) 0.00 (6) 0.00 (12)

4.0 0.00 (2) 5.00 (5) 3.57 (7)

4.4 0.00 (1) 0.00 (2) 0.00 (3)

Grand Means 4.05 (65) 22.09 (68) 13.27 (133)

 

 

57

Data for basal cell lengths from trap nests of four W
spp. are marized in Table 17. Lengths of basal cells in 60 lllll

bores were shorter for each W spp.

Table 17.

 

iiean basal cell lengths for W spp., 1986.

 

Dore depth (till!)

 

 

 

 

Wspp. 60 (N) 120 (N)
W 9.17 (3) 48.33 (3)
annulus 4.38 (21) 23.79 (28)
sunnidatus 0.00 (14) 6.37 (16)

mm: 3.72 (9) 40.25 (8)

 

 

 

58

A sill-nary of provisioned cell length data for W spp. is
presented in Table 18. These data show a possible, thoudl
inconsistent trend of decreasing cell length as bore depth increases.
The t(II) test for differences in mean cell lengths for 2 mm:
was non-significant (t = 1.7885; df - 132; p >.05, (.10). For 2.
My: differences in mean cell length for cells from 60 and 120
m bores was very significant (t - 3.2554, df = 95, p (.001).

Table 18.

 

iiean lengths of provisioned cells of W spp., 1986.

 

Hean provisioned cell lengths” (Ii)

 

Dore depths (in)

 

 

 

 

 

 

 

All t’nests 60 120
W lean (ll) liean (N) liean (ll)
J1... _ __ ._.._.
anmuam 10.8 (20) 11.10 (10) 10.50 (10)
mm 14.01 (134) 13.45 (56) 14.50 (78)
W 10.13 (96) 9.06 (36) 10.70 (61)
W 11.72 (45) 11.76 (21) 10.40 (24)

 

* The effects of occasional extraordinary cell lengths have been
cupensated for by a standard deletion of 1094 of cell length values
frail the skewed end of the data for each species.

 

 

59

Pooled data of vestibular cell lengths from W spp. trap
nests for 1986 showed differences in vestibular cell lengths were most
pronounced in bore diameter classes less than 3.2 lllll. The Z -
approximation of the liann-ilnlitney U ranks test for differences in
lengths of vestibular cells between 60 and 120 all bores was
significant (2 a 1.8826, p (.03). Vestibular cell lengths were
influenced by bore depth, with vestibular cell lengths shorter in 60
In bores in six of eidlt bore diameter classes. In 60 ill! bores with
diameters of 1.6, 2.0, and 2.4 m, vestibular cells were resectively
51.57, 42,33, and 46.42 96 shorter than vestibular cells in respective
120 ill]! bores. The mean vestibular cell length for all 60 m bores was
21.38 inn and for 120 lull bores, 34.23 all. Vestibular cell lengths in
60 lllll bores were 37.54 ’6 shorter than vestibular cells in 120
bores. Hean vestibular cell lengths for four W spp. are
given in Table 19. With the exception of E. mugging, these data
are consistent with results of the analyses of the pooled data, with

vestibular cells shorter in 60 um bores.

Table 19.

 

liean vestibular cell lengths for W spp., 1986.

 

Dore depths (inn)

 

 

 

 

W app. 60 (N) 120 (N)
111111113111: 8.50 (3) 29.67 (3)
W 23.48 (21) 38.45 (28)
gum 37.86 (14) 61.00 (16)

Wis 11.67 (9) 10.75 (8)

 

Bore Depth and Numbers of Provisioned Cells

In 1986 W produced 133 trap nests containing 363 cells

(mean = 2.77). Sixty-five trap nests of 60 lllll bore contained 157
cells (mean = 2.42). Sixty-eight trap nests of 120 iilil bore contained

206 cells (mean 3.03). Data from bore diameters of 1.6, 2.0 and 2.4

m totaled 43 trap nests of 120 m bore with 124 cells (mean 2.88) and

47 trap nests of 60 lilll bore contained 120 cells (mean 2.55).

iiean numbers of cells provisioned by W: spp. in trap

nests of two bore depth classes are given in Table 20. For three

W spp. the mean number of provisioned cells decreased in 60
bore depth trap nests coiibared to 120 m bore depth trap nests.

Statistical tests for differences in nullbers of provisioned cells per

bore depth class were significant only for 2. cumidatus.

61

Table 20.

 

liean nulber of cells per trap nest for W spp., 1986.

 

”Dore depth (m)

 

 

 

 

W spp. 60 (N) 120 (ll)
3001111111: 3.33 (3) 3.00 (3)
mm“: 2.36 (22) 2.76 (29)
W 2.62 (13) 3.70 (17)
W 2.78 (9) 3.50 (8)

 

A 100% increase in bore depth, and presumably potential volume

for nesting, resulted in an average 25.2’6 increase in nudier of

provisioned cells per trap nest. These increases were respectively by

species: 2. W, 16.95%: 2. cumidatus, 48.1%: and 2.
W, 25.9%. These results generally suggest that snail bore
trap nests with proportionately shorter bore diameters are used more
efficiently by these trap-nesting wasps. However, it is also noted
that there was no significant difference in the frequency of trap nest

selection based upon these bore depths.

Bore Opening Orientation and Frequency of Trap Nest Use

A possible concern in studies of trap-nesting bees and wasps is

whether the orientation of trap nest bore openings influence the

frequency of trap nest selection as a nesting site. The effect of

bore orientation among trap nests distributed in a forest edge was

tested by comaring orientation frequencies with trap nest selection

frequencies. Trap-nesting materials from 1984 and 1986 provided data

for this study. Trap-nesting stations were established within a mixed

hardwoods edge between a red pine plantation and an old field. Dore

openings had four orientations: south-west, facing the old field:

north-east, facing the pine woods; and north-west and south-east with

bores parallel to the long axis of the forest edge. Use of trap nests

with north-west and south-east orientations were pooled into a edm

class since field notes on those orientations were confused
Table 21 slunarizes the results of this study. The chi-square
(I) test for differences from expected frequencies of trap nest

selection shows no significant difference between south-west and

north-east orientation. Chi-square was also non-significant when

south-west, north-east and edge orientations were tested. These data

suggest that W spp. have no preference for bore orientation

among trap nests distributed in a forest edge. However, the influence

of a north-west orientation vs. a south-east orientation needs further

exami nat ion .

62

63

Table 21.

 

Bore opening orientation and frequency of use by W
spp.,1984 and 1986.

 

 

Trap nest Use
Orientation * Frequency frequency

1984 old field (S-ll) 189 18
' edge (ll-ll, S-B) 54 3

' pine forest ("-8) 189 18

1986 old field (S-ll) 360 28
" edge (ll-ll, S-E) 432 35

' pine forest (ll-B) 360 22

 

* All trap nests were placed in a mixed hardwoods edge
between a red pine plantation and an old field.

Bore Diameters and timers of Provisioned Cells

As bore diameter increased the available volume for provisioned
cells also increased but large bore trap nests did not generate more

We cells. Data from 2. My: trap nests of 1984, i985,

and 1987 were pooled to examine the effects of bore diameter on the

amber of provisioned cells. One hundred forty-six nests contained

496 cells. The nullber of cells provisioned per trap nest ranged fran

1 - 10 with a mode of 2, median of 2.85, and a mean of 3.42. Table 22

marizes data fran bore diameters 2.4 to 6.4 all. Trap nests with

bores of 3.2 - 4.8 nm accounted for 90.4 ’6 of all 2. W trap
nests and 88.9": of all 2. W cells. While there is no
significant difference in the mean nufier of cells produced in 3.2 and
4.8 II bores there is a significant difference in the numers of nests
(x2 =- 12.8444, df =- 1, p < .0005) and amber of cells (x2 =- 45.1201,
df = 1, p , .0005), prodiced in these bore classes.

 

Table 22.

65

 

Frequency of E. W trap nests and cells from 120 nun bore
trap nests from 1984, 1985, and 1987.

 

Trap nest
bore (mm)
2.4
3.2
4.0
4.8
5.6
6.4

Dore
frequency
339
609
609
609
339
369

 

cuspldatus
nests cells

9 31

61 198

42 158

28 85

2 4

3 11

 

Cells
per nest

3.44
3.25
3.76
3.04
2.00
4.67

t of
nests

 

6.21
42.07
28.97
19.31

1.38

2.07

8 of
cells

 

6.25
39.92
31.85
17.14

0.80

2.82

 

Influence of Station Species on Frequency of Trap Nest Use

Station species have received little attention in previous
research reported on trap-nesting W spp. Several factors
could contribute to the significance of station selection in
trap-nesting studies. These include the availabilty of appropriate
nesting cavities, presence of aphids, and closure materials. In 1984,
all trap-nesting stations were Jugians, and E. cuspldatus was the only
We observed and reared. For 1986, nineteen of twenty-eimt
stations were Juglans and another nine stations included 5 Enaxinus, 1
Engage, 2 Bagging, and 1 Eagus. Thirty trap nests were provisioned by
2. Mus: eldlteen by E. mmmis: six by P. annuiatus. and
fifty-one by E. W. Chi-square (I) was used to test for
differences in Eassaigegus selection of nesting stations between
Jugiang and other species. The expected frequencies were based upon
station-species frequencies: 1119.150! (67.86%) and others (32.14%).
Observed and expected frequencies of trap nest use are given in Table
23. The chi-square (I) statistic is very significant (X2 = 23.6592,
df = 2, p (.0005), and indicates that the differences in distribution
of W among these stations are not random and 2. mm
data made the strongest contribution to the chi-square statistic.

67

Table 23.

 

Station selection by three W spp., i986.

 

Juslans .chers
(laserved Expected Observed Expected

cumidatus 18 20.358 12 9.642

W 16 12.2148 2 5.7852
amiable 49 34.6086 2 12.6354

 

Data from a secondary site, consisting of stations 25 throudl
28, were particulary interesting relative to station selection. At
these four stations (three 111m and one ms) trap nest bundles
were distributed at heights of 1 to 9 meters, with 1 meter intervals.
One hundred eidlt trap nests were placed at each station, a bundle of
12 at each height. The frequencies of trap nest use at these four
stations and 9 heidlts are sun-arized in Table 24. Of 432 trap nests.
259 were used by trap-nesting wane and bees. The respective
frequencies of use at these stations were 75, 71, 52, and 61. There
was no signficant difference in the frequency of trap nest use among
these stations. However, the chi-smare (I) test for differences in
frequency of use by W was very significant (X2 = 30.03225,
df = 3, p ((.001). Fifty-five trap nests were presumed to be
provisioned by We spp. based upon closure materials, aphid

remains, and prepupae. 2. 32991111113 was found in 40 trap nests

68

collectively frml heldlts of 3, 6, and 9 m and used bore diameters of
1.6, 2.0 and 2.4 DID. 2. sum was found in 5 trap nests
collectively from heidlts of 4 and 7 ill and bore diameters of 4.0 and
4.4 lllll. While 61 trap nests were used at the 2am station by
trap-nesting wasps and bees, no trap nests at this station were used
by W. Cametition is not a likely explanation of the
exclusion of W fran the 2am: station since the arrays of
trap nest users other than W spp. among these stations were
similiar. Other factors could account for the abmnce of W
spp. from 2am. These include lack of natural nesting cavities and
closure material, such as resin, frass, and loose bark: and an absence
of aphids. These data indicate a strong correlation between station
wecies and W use. 2. My: prefers 11mins much more
strongly than does 2. W.

69

Table 24.

 

Frequency of trap nest use at secondary site by all trap-nesting
wasps and bees, and by Eassaigegua spp., 1986.

 

Height (m)

 

NCO-501014070

p

Totals

Stations and Frequency of Use

 

All Wasps and Bees

 

 

 

Juglans
9 9 4
11 10 6
5 9 1
9 9 5
10 8 5
4 5 6
9 7 8
8 7 11
10 7 6
75 71 52

Fagus

 

 

 

W spp.

Jugians Fagus
7 7 1 O
0 0 O O
0 4 0 0
7 9 2 0
0 O 0 0
0 2 0 0
3 6 6 O
0 O 1 O
0 0 O 0
17 28 10 0

 

70

Selection of trap-nesting stations for the 1987 season was based
upon the relative abundance of tree species at the study site. This
differed significantly from station selection for the Sumner 1986,
when Rings was systematically excluded as a trap-nesting station. The
species distribution of forty-nine trap-nesting stations for Sumer
1987 was Pinus - 14, Juglans - 22, Emmy: -— 8, and others (1.11m,
Assn, and Egpglus) - 5. The respective frequencies of trap nest use
by E. cuspldatus among these stations were: Ring: - 55, Juglans - 19.
[minus - 5, and others - 4 (Table 25). The chi-square (I) test
statistic is very significant for these data (X2 = 57.9884, df = 3, p

((.0005). Bing: stations are clearly preferred by E. guspidagma.

Table 25.

 

Station species fre enc and fre enc of trap nest use by
WM.1987. W Y

 

Station species

 

Pinus Juglans Fraxinus Others

 

 

 

 

Station
Frequency 14 (28.57%) 22 (44.90%) 8 (16.33%) 5 (10.20%)

Use
Frequency 55 (66.27%) 19 (22.89%) 5 (6.02%) 4 (4.82%)

 

Aphid Provisioning by 233311993113 2113215131113

E. W was selected for intensive investigation of
trap-nesting biology during the sunlner of 1987. E. W is the
largest W in the study area, its color markings (the ventral
surface of scape, dorsal surface of mandible, and basal portion of
hind tibia are white) and consistent pattern of ringing nest openings
with resin prior to provisioning, make accurate field identification
possible. Other W including W, W, and
mm: are analler and lack distinctive color markings, making
field identification uncertain.

2. W was presumably active in the study area prior to
site establishment on Nay 29, 1987. Five trap nests were ringed with
resin as of June 1, 1987. Trap nest provisioning was observed on June
1 (4 trips, 2:39 pm - 3:20 pm): June 3 (7 trips, 12:31 pm - 1:35 pm):
June 18 (16 trips, 10:17 am - 11:43 am); and July 5 (8 trips, 4:31 pm
to 5:13 pm). Nesting activities continued througl the first week of
Auwst with no trap nests ringed or closed with resin after August 6,
1987. Cessation of E. cuspldatus activity was verified by
observations on August 8, 14, and 21, 1987, which provided no evidence
of additional activity. The minimum provisioning period for this was)
population was seventy days. This corresponds well with the range of

flimt dates (June 14 - August 14) for E. cuspldatus material from the

71

72

Museum of Zoology of the University of Michigan and the Entomology
Museum of Michigan ‘State University, and is supported by my
observations of E. cuspldatus activity in 1984 and 1985. in 1984,
provisioning activity was first observed between June 4 and 12 and
terminated between August 4 and 13. In 1985, emergence of E.
cumidatus from a natural nest occurred May 10 and nesting activity
ceased August 20.

The provisioning period of 2. W for 1987 was divided
into seven, 10-day intervals and tallies were made of ringed and
closed trap nests for each interval and are given in Figures 8 and 9.
Increasing nunbers of closed trap nests in intervals three and four
suggest that E. W made a behavioral response to increased
numbers of prey. The chi-square (I) test for differences in closure
frequency for these intervals is significant (X2 = 24.8387, df = 6, p
<.001).

equencu of Ringed and Closed Iran as
by Passaioecus cuspldatus, 1387.

 

 

 

 

Ten-dag intervals of‘ Trapnesting Season
.Ringed Trap Nests .Ciosed Trap Nests

e e cy of ringed and closed trap nests during 10—day
ls nnin nd ending August 6, i987.

CODI'DCQI‘D'S“

Figure 9.

74

Cummulative Fr equenog ol‘ Ringed and Closed
Trap Nests b9 Passaloecus cuspldatus, 1987.

 

V25-Vl1
VIC-17
WHO-27
V128 Vll7
Vll8-17
WHO 21
VII 28-Vlll8

Ten-dag intervals of‘ Trapnesting Season
.Ringed Trap Nests fiClosed Trap Nests

Cummulative frequencies of ringed and closed trap nests
during iO-day intervals beginning May 29 and ending
August 6, 1987.

75

E. cumidatus used Cinaria sp., W 59.. Elm-nnin: 89..
mm sunbathing (Thomas), 11121.1: sp., Mugs ms; (Fabricius),
111m W (Davis), and 51mm mung Fabricius as provisions.
Table 26 stmIIiarizes data on six presumed types of aphid provisions.
Multiple t(II) tests for differences in nunber of aphids provisioned
are significant for Mm: W, 9.1112213 sp., and MW
munching, [flung mum and 91mm sp., t(II) = 7.05053, df =
193, p (.001: mm Sp. and W W , t(II) -=

3.62055. df = 72. p < .001: and Mimi: mnardae and Macrosiehlml
mm, t(II) =- 10.7478, df = 197, p < .0011. Differences in

number of aphids provisioned can be correlated to relative aphid size,
with any: m the anallest and Wm W the
largest. In some cases larval development in interior cells was quite
advanced and much of the provisons were consumed. Thus the mean
nunber of aphids provisioned per cell is a minimum, the true mean

being slidltly higher for each species of aphid provisioned.

Table 26.

 

Aphids provisioned by 2. W, 1987.

 

lean
A hid Number of Number of * of Standard Standard
ype* Trapnests Cells Aphids Deviation Error

1 46 160 37.0375 11.8080 .9335
2 9 35 26.8287 7.6625 1.2952
3 12 39 21.7949 8.2902 1.4882
4 3 8 28.5000 7.7298 4.1748
5 3 8 43.0000 17.5770 2.7329
6 1 6 34.8333 9.0077 3.6774

 

*1= 12=Cinaria . 3=
WWW sp. and’fixéu: 89.; = 122L531:
- amas-

76

The range of numbers of aphids provisioned per cell for aphid
species 1 throudl 6 were: 9 - 74, 13 - 51, 9 - 43, 18 - 39, 23 - 74.
and 24 — 47 respectively. The mean number of aphids provisioned per
trap nest were: species 1 - 128.83: species 2 - 104.33: and species 3
- 70.75. The mean numnr of cells provisioned by E. cuspldatus for
these three aphid species were: species 1 - 3.48: species 2 - 3.89:
and species 3 - 3.25.

Two trap nests had an extraordinary manner of provisions. One
contained 430 aphids (eight cells, 53.75 aphids per cell) and another
contained 334 aphids (five cells, 66.80 aphids per cell).
Unfortunately, aphid samples for identification were not taken from
these trap nests.

Figure 10 summarizes data on the seasonal changes in aphid
selection by E. W and relative numbers of aphids
provisioned. B. W was not restricted to a particular aphid
species and was able to switch to available resources. Peaks in
provisioning rates were assumed to be related to increased aphid
numbers. Figures 11 and 12 summarize data on aphid provisioning
activities. The provisioning period was divided into fourteen 5-day
intervals. The number of aphids provisioned in each interval were
tallied. initial provisioning activity was moderate and five trap
nests were ringed with resin within one or two days of trap nest
station establishment. Between May 29 and June 7, man sp.,
m1: sp., My: sp., and 51mm am Fabricus were

.hmou ups-35m .33 .N

am nouuo_oo soon :. noncocu .ocoooom .9" 09:0.h

H'LIIM
S-aflM
HIM-8311A

53am 22:333.. 5 32:35

LL-Ci llA
Zl-IIIA
8

' ZIM'UZM
l” R'IHA

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

.... .13;le 1» \KJ
...... a CR / N
383a 41W / u
8.1.3 s (\

 

 

32 555.5
carom—am as: 333.250 33333..

 

[eniaiul / pauoislnoid splqdu JO iaqulnN

78

.FOOu .@ u0303¢ Oc-Uco 0:0 ON >02 Oc_cc_mon n_a>90u=— >flUIm

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

3.38mi. ... .3 commmm mflEomeWE c” ...;MEEAE A
Egg w. .w \: 1....
§ mm
E§ “up
\ as:
g “a
ge—

 

 

 

 

 

 

.m_m?.35 33-3: 5325... mars:
3.3325 .n. as 35339:. 32.5 .3 33:52

a: 09-- C n_::---'U In

79

.53 6 539% 9:93 new on >2. 9:533 2233.: >mvum 9:26
nan—wanna .m 3 3:30:63 32% no .395: 3322550 .Nd 9.3:

33333 ... .3 23% 22°33; E 22:35

 

 

 

 

 

3: 0&— C: u-I—U u'l

 

 

 

 

 

£2.35 :25»: 5332 9:3: mBmEamao ...
ma 3:223; 2.2:: .3 33:52 3:22:58

80
used as provisions. From June 8 through July 27, My: W
(Davis) was the preferred prey while Wm W (Thomas)
and may; gems], (Fabricius) were provisioned in significant numbers
between June 28 and July 17.

Provisioning rates were quite varied and were estimated on the
basis of number of cells provisioned and dates of bore ringing and
closure. A scatter-gram (Figure 13. A.) summarizes data from 79 trap
nests. Trap nests with estimated provisioning periods of one to three
days contained from one to ten provisioned cells. Trap nests with
estimated provisioning periods of 6 to 8 days contained two to eidit
provisioned cells. Trap nests with provisioning periods of 10 to 21
days contained one to five provisioned cells. The estimated number of
cells provisioned per day ranged from a low of 0.1 to a high of 5.
Several variables could influence provisioning rates. Exceptionally
high provisioning rates could be the result of close proximity of
aphids and closure materials to the nesting station. Aphids and
closure materials sanewhat removed from the nesting station could
produce low provisioning rates. An additional possible cause for low
provisioning rates would be a temorary cessation of provisioning
activity. This cessation might be necessary to allow the development
of additional ova following a period of provisioning and oviposition
activity. Two hundred eighty cells were provisioned during a
cummulative provisioning period of 381 days. The average provisioning
rate per trap nest was .7349 cells per day. Trap nests worked less
than four days had a higher than average provisioning rate and trap
nests worked longer than four days had lower than average provisioning

rates.

81

Least squares sinple linear regression of average numbers of
cells provisioned and length of the provisioning period per trap nest
produced the regression equation Y = -.20X + 3.99 (Figure 13. B.) and
a correlation coefficient of -.59, suggesting a weak negative
relationship between length of the provisioning period and nunber of
cells provisioned. ivlith the deletion of five extreme data points
(Figure 13.: m, n. (J . p, and q), least squares regression produced
the equation Y a .30X + 2.12 (Figure 13. C.) and a correlation
coefficient of .78. indicating a moderate positive correlation between
the duration of a trap nest provisioning period and the nunber of

cells provisioned.

62

11- A
10- -"‘

U n

E"

:6»: 8‘ ‘

E7- 0 u

a

2 .o

'76

o

'5

4::

 

 

 

III I r: l ["7"
7891011|22|

1:..4
(n
0)

I I
1 2 3

Trap nest provisioning period (days)

Figure 13. Provisioned cell prodnction rates for E. W.

Economics of Cell Partitions and Closures

Resin gathering activity was oburved at resin flows on Elm
m on June i7. 198?. One and one-half inch fence staples had
been used to secure trap nest bundle carriers to trunks of trees
selected as trap-nesting stations. Resin flows were promced in
response to fence staple wounds. 2. My: selected resin flows
with dimensions 9 In by 4 m. The wasp’s mandibles were used in a
scissors-like fashion to excise a drop of resin with a diameter the
width of the waep’s head. when separation of a resin drop was nearly
cmiete, the was: backed directly away from the resin flow. A thin
strand of resin, connecting the resin drop to the flow. was drawn into
the excised drop by lateral and circular motions of the wasp’s head.
Any remaining remnant of the resin strand was cut off by a continued
backward movement entwined with an abrubt turning to the left or
rid'lt. Mnty-two resin gathering trips were observed between 9:47 am
and 11:42 am. Resin drops were carried on the ventral surface of the
mandibles.

Three separate resin flows were used daring these resin gathering
activities. Resin flow (I) was visited repeatedly and the was:
returned directly to the resin flow. landing within a few centimeters
of the flow and approaching it directly. After a nutter of resin
gathering trips, the remaining portion of the resin flow was too
sail, or of inproper consistency. and was abandoned as a resin

“FCC .

84

The 21m trunk was searched for another appropriate resin flow
and resin gathering resumed. Resin flow (II) was then used repeatedly
as a resin source. After three or four trips to resin flow (Ii) the
wasp made a trip to resin flow (I), explored the resin mass. and
returned to resin gathering at flow (11). Resin gathering was again
observed June 18 (eimt trips. 4:31 pm - 5:13 pm) and July 24 (11
trips between 9:00 am and 10:00 am. with no elapsed time recorded).
Ringing of trap nest bore openings was also observed and the following
action pattern was noted. 2. We landed at or within a few
centimeters of the trap nest opening, entered head first, exited and
re-entered gaster first. The wasp then appeared at the nest opening
and with her mandibles unread a thin layer of resin on the face of
trap nest at the margin of the opening.

Kronbein (1967) noted that resin partitions were umaily .25 and
occasionally 4.0 m thick. while closures, ranging frm .25 to 4.0 an.
were usually 1.0 III thick. Vincent (1978) noted that 4 drops of resin
were used for nest closures. Data from my studies of 1987 were
examined to determine the variability of partition and closure
dimensions, their volumes. and the energetics of resin gathering. It
was noted curing resin gathering described above that an excised drop
of resin had a diameter approximately equal to the width of the waqz’s
head. Given a diameter of 1.5 In. a resin drop carried by E.
W has a volume of 1.77 m3. The thickness of resin
partitions of 295 provisioned cells ranged from 0.1 to 5.0 m in trap
nests with bore diameters 2.4 to 6.4 nm. Median and modal partition

thicknesses were 0.5 In; the mean partition thickness was 0.66 m.

85

Seventy-one closures had thicknesses ranging from 0.25 to 4.0 m with
a mean of 1.69 cm.
Based upon the bore diameters and thicknesses indicated above, the

volume of resin required for partitions and closures ranged from 1.13

m3 to 62.83 m3. The volumes of resin required for partitions and
closures in bores most frequently used by E. W are
respectively: 3.2 an - 5.31 and 13.59: 4.0 an - 8.29 and 21.24: and
4.8 l- - 11.94 and 30.58 m3. Given a volume of 1.77 m3 per resin
drop. the nmrs of resin gathering trips for partitions and
closures for these respective bores are: 3.2 an - 3 and 7.68 (8): 4.0
II - 4.68 (5) and 12.0: 4.8 nu - 6.75 (7) and 17.28 (18). These data
are rather conservative since they have not taken into consideration
the foundations for closures or partitions. Foundations. consisting
of a resin ring on the wall of the trap nest bore, have base widths
greater than that of their respective partition or closure. Resin
partitions, closures, and their respective foundations, represent a
significant energy investment. Resin volumes and bore diameters
across which resin must be drawn possibly contribute to the upper
limits of the bore diameters used by E. W. The distribution
of partition and closure thicknesses frail 2. gym trap nests are

given in Fiwre 14.

 

O. 0
use: 9.: «33398 a 5 2:320 23 26332:. 508 «o 32:25.

Op

255 3252.:
n v a N p D. v. a.

Lari}.

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

32535 3:38-

 

 

....3525 52:53

 

 

 

 

 

 

 

 

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ON
an

0v

ON

on

con
0:
ON—
on—

ovp

.vq annu—m

msmrsucom

SUNHARY AND CONCLUSIONS

Several W spp. were found in a snail study area on the
camus of Concordia College. Ann Arbor, lilchigan. 213311.022“: spp.
that used artificial nesting materials in the area included: 2.
W. E. W. E. cumidatus. P.- umilJmnan. and B-
M. 2. um: and 2. mm were also collected fran the
cupus area but were not reared from trap nests at the study site.

Trap nests with bores greater than 4.8 m were rarely used by
W spp. while diameters of 1.6 - 4.8 m gave good results.
We nesting sites were partitioned on the basis of bore
diueter, height. and station species. W cumidatus
preferred Emu; stations. This preference is possibly associated with
the availability of resin, which was enhanced by the method used to
secure trap nest bundles to trap-nesting stations. 2. W also
preferred trap nests heldxts below 3 meters and bore diameters fral
2.0 to 4.8 nm. W mm preferred Juglans stations. trap
nest heimts above 3 meters and bore diameters 1.6 - 2.4 an. Bore
selection seems to be based upon wasp size while heidit distribution
could be influenced by the usual distribution of natural nest sites.
It midit be suggested that larger diameter natural nesting sites are
distributed at lower heights and saaller natural nesting sites at
greater heights.

Trap nest architecture. including the lengths of basal cells and
vestibular cells was influenced by bore diameters and bore depths. An

87

inverse relationship was found between bore diameter and cell length.
Increased bore diameters resulted in decreased lengths of provisioned
cells. while activities of parasites were associated with increased
cell lengths. wasp senescence did not result in increased lengths of
provisioned cells. Volumes of provisioned cells increased along with
increased bore diameter, but numbers of aphids provisioned did not
increase proportionately with increased volume. Bore vodume was not
maximally used for provisioned cells. Basal cell lengths. in trap
nests with bore diameters less than 3.2 mm, were signficantiy reduced
in trap nests of 60 mm bore depths. The number of provisioned cells
per trap nest increased for all Egssglgggug spp. as bore depth
increased, but was not statistically significant. hall bore trap
nests with 60 Ill! depths were used more efficiently that trap nests
with 120 n depths.

Easaalgegus cuspldatus were active from late May through early
Aumst. and provisioned at eidut species of aphids. The nunbers of
aphids provisioned differed significantly among three aphid species
and was inversely related to the size of the aphid. Two hundred
eighty cells were provisioned during 381 provisioning days for a rate
of .7349 cells per day. The nunbers of aphids used as provisions
varied dramatically (9 - 74). While snail variations could be
explained on the basis of differences in sizes of aphids. such large
differences are more problematical. Suggested causes of large
variation include sudden weather changes and the disruptive behavior

of parasites.

89

W spp. were parasitized by m: ms, Q.
W.mmm.mmmm.andmm
slum.

These studies add significantly to the literature on the biology
of Eassaigggus. The efficiency of small bores (1.6 - 4.8 mm) and bore
depths of 60 nm is demonstrated for snail trap-nesting species.
Techniques and methods described in this paper can be applied to the
aphid provisioning activity of other W. This study also
shows that Eassalgggus should be added to the classical list of aphid
predators. Finally, artificial nesting materials were used very
successfully and provided an inexpensive and biologically interesting

technique for the study of aphid hunting wasps.

APPENDICES

90
APPENDIX 1

Record of Deposition of Voucher Specimens*

The specimens listed on the following sheet(s) have been deposited in
the named museum(s) as samples of those species or other taxs which were
used in this research. Voucher recognition labels bearing the Voucher
No. have been attached or included in fluid-preserved specimens.

Voucher No.: 1989-08

Title of thesis or dissertation (or other research projects):

TRAP—NESTING BIOLOGY OF PASSALOECUS CUSPIDATUS SMITH (HYMENEOPTERA:
SPHECIDAE) AND SYMPATRIC SPECIES

 

Museum(s) where deposited and abbreviations for table on following sheets:
Entomology Museum, Michigan State University (MSU)

Other Museums:

Investigator's Name (3) (typed)
John Morris Fricke

 

 

Date October 10. 1989

*Reference: Yoshimoto, C. M. 1978. Voucher Specimens for Entomology in
North America. Bull. Entomol. Soc. Amer. 24:141-42.

Deposit as follows:

Original: Include as Appendix 1 in ribbon copy of thesis or
dissertation.

Copies: Included as Appendix 1 in copies of thesis or dissertation.
Museum(s) files.
Research project files.

This form is available from and the Voucher No. is assigned by the Curator,
Michigan State University Entomology Museum.

91
APPENDIX 1.1
Vbucher Specimen Data

Pages

12

of

l

Page

as same 2%

huanue>uca oumum sewage“: one as uwmoees

mama

i nousrmDWV

 

$3 -3 $0.88 38

 

 

 

you msuawueem seumaH u>onm use so>wuuum

 

melmmma

.oz menuso>

Aeuasav

luquum mamwmmlmnmdi

Amvusmz m.uoumwwumu>sH

Amusmmooos aw mumo£m Hmcoauwvvs umav

 

3m:
3m:

3m:

3m:

Nimmimmiam smmcomse
Hamoiomiem smmcemse
same .c~ 2m:
exodus .2 snow

.00 swamucmmz

posse ace ":UH2

snimmim sameness
exudes .2 ages
smms .52 2m2

.ou 3mcmunmm3
gonna :24 "mqu

omelmmim sameness
excess .2 anon
pmma .m2 2M2

.00 3mcwu£mm3
scene 2:2 "onz

 

A>mmv mzumazccm msooonmmmm

 

A>mmv mzumasccm msumonmmmm

 

A>mmv msumasccm msumonmmmm

 

Museum
where

depos-
ited

Other

 

 

Adults 0v

 

 

Adults 9
Pupae

 

 

Larvae
Eggs

 

vouumoaus vsm sums no souomaaou
momEHuoam pew mums Honda

 

:oxmu uenuo mo mmaoeem

 

 

"mo sunfisz

 

 

92

APPENDIX 1.1

Vbucher Specimen Data

Page 2 of

12 Pages

new same §§.§

huwmuo>uc= oumum damaged: any as uumonuv

mama

HON ”MB

 

mmma 0H HBOHUO mama

 

 

 

now msoauooem emumwu o>oam ozu ve>wuuu¢

 

menses mmuuoz anon

 

 

 

 

 

melmmma .oz uuzoso> Asmezuv Amvuamz m.u0usw«umu>su

Aaummmeuos ma masons Hecoauuvss emav
2m2 H Humniomumm unmadmue
:m2 H Nipmuomrmm sameness
:m2 a Himmiomuam “message
am: A wimhlmml<m nauseous
3m: H Hiahlmmlem escapees
am: A Hiahlomlma ummcomua
mxofium .2 snow
hams .mm >m2
.ou encounmms

Henna sac ”mqu acmocfl> msumaooum msomonmmmm
am: A museummlmm ummcomue
amz a oiamimmicm sameness
am: A mlaolomlsm ummcamue
smz H miaoimmimw sameness
3m: H Nlmolmmlma assesses
exoaum .2 anon
smma .m2 2m2
.ou encounmmz

nouns use "30H: A>mmv manganese msumonmmmm

zoo? souqmomos use sums no seuumuaoo :oxmu nonuo no mowumqm
m e r r m m e .m u msuauoeam you muss Honda
mmmmmmmmm;
“M w.d.i nu in in p. n“ r“ we

 

 

 

 

 

 

 

 

 

“mo amassz

 

 

93
APPENDIX 1.1
voucher Specimen Data

Page 3 of 12 Pages

sesame res,

huamuo>aca eumum smwwnuuz any :« uumoeas

muse

#09 9

 

$3 3 H338 33

 

 

IQ

 

ems: honoEoucm

 

pow m:eE«omem soumwfl m>ons msu so>auuum

 

mOImmmH .oz umnuso>

mxoaum mfluuoz snow

Avonhuv Amvmamz m.uouquumm>cu

.A2ummmuoec ma mousse Hmsouuwsvm mmav

 

3m:
3m:
3m:

3m:

3m:
3m:
3m:

Firlrl

P4F1F4

Hiamimmlaa sameness
Nivpuomlaa ummcamue
Nummlomlas ummcamse
exodus .2 snow

peas .oN >m2

.OU 3mcmu£mm3

posse ace "on2

Nimnimmiam awesompe
exodus .2 anon

hams .Hm >m2

.ou Secounmmz

gonna ace "mon

mimoimmiam “message
mimolmmram smwcomse
Himoammiam sameness
excess .2 econ

hams .mw >m2

.OU swamunmmz

gonna ace "20H:

 

ucmoca> maumaoesm mausonmmmm

 

UCQUCH> MDHMHOmhm m30¢0H0mmmm

 

ucmoca> mzumaomum msomonmmmm

 

Museum
where

depos-
ited

 

Other

Adults 0'

 

S
h
p

m

Adults 9
Pupae

 

 

 

 

Larvae
Eggs

 

vouumoaev was see: no vouueafioo
mauEaueam MOM mums Hens;

 

:oxmu pesuo no mouuenm

 

 

uuo sunfisz

 

 

94

APPENDIX 1.1
Voucher Specimen Data

Pages

12

of

_:L_.

Page

 

mama .OH “manage mum:

 

 

 

mm? «‘me ruwoufiijnw 5w.

a: honoEOucm

 

huumuu>uc= wumum cmmusuaz usu :« uumonum
uou mcmefiuoam muumwm u>onm «nu vu>uuuu¢

 

meHum.mwuuoz anon

monmmma .oz uazuao> AcmcauV Amqumz m.uoumm«umm>cn

Azummmuuu: mu mucosa Hmcoauuvvu omav

 

3m:

3m:

3m:
am:
3m:
3m:
3m:
3m:
3m:

mxuflum .2 2202
q hmma .OH >m2

.00 3mcmu£mmz
conga ace "muH2

mxofium .2 anon
o hmma .q 2m2

.00 3mcmu£mm3
gonna see ”20H2

vuamummu<m ummcamue
muamuwmnam ummcamue
mIMhuomuem umwcomue
mumnuomucm ummcamue
enmhuomuam ummcomue
Nuanuomuma ummcomue
Huanummnma ummcomue
mxoflum .2 anon

pmaa .om >m2

. 0U 3mcwu£mm3

gonna :22 "2UH2

HHHHHHI—I

 

:uHEm msumodomso msumonmmmm

 

cufiem msumuflommw mzomonmmmm

 

ucmocfi> maumaowum msumonmmmm

 

Museum
where

depos-
ited

 

Other

vmuuwoaov mam mom: no mmuowdfiou
mausaomam he“ mums Hanna

 

 

 

 

 

 

Adults 0‘
Larvae

Pupae
Nymphs
Eggs

 

coxmu mango no muuuwam

 

 

“5 Adults 9

umaeaz

 

 

APPENDIX 1.1
Voucher Specimen Data

Page 5 of 12 Pages

2&0

«mm:

.20 w Qua. 9w?

zuamuo>20= oumum oowuzouz on» 0d uuuoooo

HO“ whfllok.

 

mmmH om um20uoo mug;

 

 

 

you m0oefiooom vouowa o>ono ozu uo>wooom

 

mxomum mflummaumaamw

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

melmmma .oz nozooo> Aooazuv Amvusmz m.uoumw«umo>0a
Axummmuooc mu mucosa Hooouuuovo «may
020900 .2 0000
:m2 m mmma .mm >222
.00 zmcmunmmz
nonuc 00¢ "30H2 eonanma mmumasvcmm mdumOHmmmmm
mxuaum .2 0:00
:m2 UH mmma .m mm2
.00 zmcmucmmz EODHLmo
nonu< 00¢ "20H2 mfl0u00mam005 msomoaommmm
mxomum .2 0000
2m2 om mmma .ca >m2
.00 300000003 eonacmo
Honuc 00¢ ":0H2 mm0u00HHa0oe msomOHommmm
mxomum .2 0:00
2m2 om mmmm .m >m2
.00 3m0mu£mmz
nonum 2:4 "20H2 sumem mammomomsu mnemoaommmm
100+ vouamoaoc 000 mom: uo mouoogaoo ooxmu nosuo no moaooam
m e r r m m e .M a m005uooom you mumm #0004
e r 0.0 e .1 .1 a w s
mummmmmwm.m
deiOAAPNLrWI :

"no M00502

 

 

 

 

mum: gowwgom

 

mama 0H MOQOpUO mama

 

 

N9 4.3%; i... Mugflfi

 

>uugg0>q0= ououm snagged: «so 0“ uqmoaoo
gou moosvuoom vogmma o>ooo ozu mm>wooox

 

meAgm magg02 0:00

monmmma .oz gozooo> 2000200 Amvuemz m.goumm«gmo>0u

Aagmmoooo0 «a mucosa Hooowugmvm 00:0

 

Pages

of 12

6

APPENDIX 1.1
Voucher Specimen Data

Page

am:

2m:

3m:

oxoggm .2 once
m mmmg .mg ooze

.OU 3mcwu£mm3
googa com "20g2

oxoggm .2 once
N Nmmg .o~ mgso

.00 3mcmu£mmz
gonga com "20g2

oxoggm .2 anon
m ommg .mg mm2

.OU 3mc0u£mm3
googm com "20g2

 

sonanma magmaomcfim m0000Hmmmmm

 

eonanmo magmHDUCmm mSUOOHmmmmm

 

sonanma magmaso0mm m5000ammmmm

 

Museum

where
depos-
ited

 

Other

voummoaom 000 ton: go mogoofimoo
mcoEuooam gou mums doom;

 

 

 

 

 

 

Adults 0'
Adults 9
Pupae
Nymphs
Larvae
jEggs

 

ooxmg gosuo go mofiooam

 

 

o Monasz

O.
“-1

 

 

9?
APPENDIX 1.1

Voucher Specimen Data

ges

of 12 Pa

21...

Page

 

 

mmma 12 gonoooo 38

 

 

 

19 .wwmt goumuoo
S. .020 Sfiflwfim

 

>uuug~>w0= oumum oom«:o«x 0:» 0a gunooom
gou moosqoono oouo«m o>ono o:u mo>uoouz

 

mxomgm mmgg02 0:00

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

woummma, .oz gosooo> Ammomuv Amvmsmz m.gogmw«umo>0u
Ahummmfluufl u.“ DUUOSQ HQCOdUfiU—ud 02.:
:m2 “Anna g 0mm gmocomge
am2 gmoom g mam Homepage
:m2 gmooc g oog gmooomge
2m2 gmool H mm gmocomge
82 332 N mg gmocooge
Dm2 mflmscm N moo ummcamge
oxoggm .2 coon
mmmg .mg-m >m2
.00 zmcousmmz
uOflud 50¢ "IUHS Amflflofluflmmv WDGCQG gHmEO
oxoggm .2 2202
:m2 g mmmg .om gmoooa
:m2 N mmmg .g gooogoo
2m2 g mmmg .mm mgoo
am2 g Nmmg .o gmoosa
.OU 3mcmu£mm3
.uOQHd rad ”IUHE EODHLmQ ESCOfiHDu. MSUOOHmmmmm
400+ vouqmoaom 000 mom: go mouoouaoo coxmu gmsuo no mowooam
m e r r m m e .m a moosqooom now 0000 #0004
e r 0 d 2 1m 1 a W 8
e D. e h u D. 8
mm. mm m g n m m m ....
“no gmasoz

 

mmmg .og gonogoo oggo

 

 

 

fawo~oaoucm

>0u000>uca 0u0um 00mm00q= 000 0a uuuoaov
gou 000000000 00uoua 0>o00 00a 00>«0000
moummma. .oz g00o=o> 2000200 A00050z 0.0og0wuum0>0u

A>u0mmoo00 um 000000 H00ouummc0 000v

 

020Hum magg02 0000

 

 

A we mmumm 000000g9
NI Hmlomlmo 00000009
m vmuomumm “mocomge
H mm mmaam gmocomge
oxoggm .2 snow

mmmg .m 202

.00 300000003

g00u¢ 00¢ ":0H2 200000000 m0owoam 0H00500

3m:
3m:
3m:
3m:

Fir-UHF!

 

Dm2 01001 a ma um0000g9
oxomgm .2 0000
mmmg .m moon

APPENDIX 1.1
Voucher Specimen Data

Page 3 of 12 Pages

3m:

.00 300000003
HODM¢ 00¢ "30H:

mm 000000gs
0xomgm .2 0000
mmmg .m mono
.00 300000003
gonu¢ 00¢ "00H:

 

Amsmoflgommv 000000 madmam

 

20000000000 000000 msameo

 

Museum
where
depos-
ited

 

Other

 

 

 

 

 

Pupae
Nymphs

Larvae
Eggs

 

000000000 000 000: 00 m0uo0~Hoo
000500000 you 0000 H0000

 

00x00 u00uo no mouoonm

 

 

“005:2

 

 

. 0.0.2.30 m -.....m

 

 

mmmg .og gonogoo oggm

 

 

 

 

 

0:2 mmoa
huauuo>aca 000um 00wu00q2 00y 0m um00000 “WMUHHHanflnaflAHfimfl.
you 000300000 0000«H 0>o00 00a 00>«0000 . .
moummma .oz u00ooo> A000>uv Amv0a0z m.uou0w«um0>0u

A200000000 «a 000000 0000002000 00=v

 

Pages

of 12

APPENDIX 1.1
Voucher Specimen Data

9

Page

Dm2
3m:

3m:
3m:
3m:
3m:

H mamonomuaa ammoomge
a vuqmummumg gmooooge
oxoggm .2 econ
mmmg .00 >02
.00 300000003
g00g¢ 00¢ ":0H2 A00000g00 000H0H0 0000500

 

guamnomnmm gmocomge
muomuomumm “monomge
mumm-omuam umocomge
mImmummuaa gmocomge
oxofigm .2 anon

gmma .m >m2

. CU 3mcmu£wm3

g00g¢ 00¢ ":0H2 200000g00 m0ofiDH0 0000600

r-{I-IHv-I

 

 

Museum
where

depos-
ited

00uumon00 000 000: no 00000HH00 00x00 ~00uo no 00uo0nm
000emo000 gou 0000 #0000

 

 

 

 

 

 

 

 

Adults 0'
Adults 9
Larvae
Eggs

Other
Pupae

 

"mo 000502

 

 

 

100
APPENDIX 1.1

Voucher Specimen Data

Page 19 of 12 Pages

000000.: . .

0% {3... m $0....NV... 230$

 

000000>00= 0000m 00000002 000 00 0000000

000:

hOukfih—HD

 

mmma .OH 0000000 0000

 

 

.500002 0m000500cm

 

000 000500000 000000 0>o00 000 00>«0000

 

monmmma

.02 000000>

A000>0v

.000000 mflggo2 snow

Amv050z 0.000000000>0H

2000000000 00 000000 H000fi0u000 00:0

 

3m:

3m:

2m:

3m:

mmo-mm-~ 00000000

00050 0000000000 00000000000
0200g0 .2 anon

.00 20:00:00: .0o0g< 20< "0002

mmo-mm-~ 00000000

00050 0000000000 00000000000
0200g0 .2 snow

.00 300000003 .gonum :00 ”0002

oHHImmIH 000000mw

00050 0000000000 00000H00000
0200g0 .2 snow

.00 300002002 .00002 :04 "0002

I 01H 00000009
00050 000000W000 00000000000

000000 .2 0000
.00 300000002 .0000¢ 00¢ "00H2

 

 

00050 0000000000 00000H00000

 

Museum
where

depos-
ited

 

Other 0

 

Adults 6'

 

 

 

 

Pupae
Nymphs

Larvae
Eggs

 

000000000 000 0000 00 000000000

000500000 000 0000 00000

 

00x00 00000 00 0000000

 

 

“5 Adults 9

000502

 

 

101

APPENDIX 1.1
VOucher Specimen Data

of 12 Pages

1

Page

030020 . .

 

000:

20.000 50.0.0 03wa

>00000>000 00000 0000000: 000 00 0000000

.ImmmdliqdluuaaHUQI 0000

 

 

00: 0000050000

 

000 000500000 000000 0>000 000 00>0000¢

 

molmmmH .oz 000000>

A000>0v

 

000000 000002 0000

Amvuemz 0.000000000>0H

0000000000 00 000000 0000000000 0000

 

302

302

302

302

maelmmum 00000009

00050 0000000000 00000000000

. 000000 .2 0000

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102

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APPENDIX 2

Preliminary Surveys and Selection of Study Area

During the summers of 1982 and 1983 malaise trap (Towne’s design)
collections were made to survey in general the Hymenoptera of
Concordia College, Ann Arbor. Michigan. The 235 acre campus is the
former Earhart Estate. Pine plantations, mixed hardwoods, old fields
and meadows provide generous areas for study removed from the central
campus. Table 27 and Table 28 respectively give the families of
Hymenoptera and sub-families and tribes of the Sphecidae that were
collected. ‘The Eassalgegus, collected included the following: 2,
cumidatus Smith - 3: E. uneasy: Vincent - 2: 2. WELL: Dahlbom -
42; and E. tunignum,Dahlbom - 4.

103

104

Table 27

 

 

Families of Hymenoptera collected at Concordia College,
Ann Arbor. Michigan, during the Summers of 1982 and 1983
using a malaise trap of Towne’s design.

 

Pamphiliidae Euchartidae Diaprildae Eumenidae
Argidae Perilamphidae Scleionidae Pompllldae
Diprlonldae Torymldae Chrysidldae Sphecidae
Tenthridinidae Pteromal idae Bethyl idae Col let idae
Sirlcidae Chalcididae Dryinidae Halictldae
lehldriidae Cynipldae Tiphiidae Andrenldae
Braconidae Gasteruptlidae Mutlliidae Hegachilldae
Ichneumonidae Helorldae Pormicldae Anthophorldae
Hymaridae Proctotrupldae Vespidae Apldae

Eulophidae Ceraphronldae

 

105
Table 28

 

Sub-families and tribes of the Sphecidae collected at
Concordia College, Ann Arbor, Michigan, during the summers
of 1982 and 1983 using a malaise trap of Towne’s design.

 

Spheclnae Larrlnae Nyssoninae
Spheclni Larrinl Alyssonlni
Scelephronlni Trypoxylonlni Nyssonlni

Hiscophlni Gorytini

Astatinae Crabronlnae Phllanthlnae

Astatlni Crabronini Philanthlni
Cercerinl

Pemphredonlnae
Psenlnl
Pemphredonini

 

Concurrent with malaise trap collecting. trap-nesting studies
were conducted between June 30. l982, and August 26. 1983, to
determine the effectiveness of trap-nesting techniques and the
presence of trap-nesting wasps, especially We. Trap nests
were constructed using techniques described by Krombeln (196?). Trap
nest bundles were assembled for field distribution using Pye’s methods
(Fye, 1965). For the 1982 season trap nests consisted of clear white
pine blocks (19 X 19 X 152 mm) drilled to a depth of 114 mm. Bore
diameters were nominally 3.2, 4.8, 6.4, 8.0, and 9.6 nm. Trap nests
were bound together into 3 X 3 bundles of nine trap nests. Bundles
were attached to 2’ X 2’ wooden stakes, four bundles to a stake.
Three sites were selected and three stakes (4 bundles of 9 trap nests
each) were placed at each site (Figure 15). Site A was a forest edge
addacent to the campus ponds; Site B was a wooded fence row southwest
of St. Paul Lutheran School; and Site C was an edge between a pine

plantation and an old field that faces the Huron River.

106

 

 

Huron River

Figure 15. Schematic map of Concordia College. Ann Arbor. Hichigan.
indicating trap-nesting sites for 1982.

107

Three hundred twenty-four pine trap nests were placed in the
field. In late September these materials were removed from the field
sites. One stake from Site A and another from Site B were lost due to
vandalism. Between October 4, 1983, and March 18. 1983, the remaining
252 trap nests were opened and 40 contained overwintering contents.
Larvae and prepupae were removed from some trap nests and stored in
alcohol vials. Others were placed in rearing tubes to rear out
adults. Adults of the following families were obtained: Bombyllldae.
Braconidae, Chrysididae, Eumenldae, Ichneumonidae, and Sphecidae
(Sphecinae: Isodontla: Larrlnae: Trypoxylonlni). No Easaalgegga were
reared out.

Trap-nesting studies for the 1983 season were expanded in number
of nests, sites. and variety of nesting materials. Nesting materials
included Rugs twigs, 13 - 25 mm X 150 mm: One-to-two year old Enaxinus
twigs. 150 m long: plastic straws, 165 um long with a 3.6 nm bore:
and pine trap nests. Between May 3. i983, and June 20. 1983, five
sites were established (Figure 16).

Site I was near the central campus. One hundred forty-four pine
trap nests were placed adJacent to a cedar border between the
greenhouse and Manor 1. Each of four stations at this site consisted
of four 3 X 3 bundles on wooden stakes. A E. W female was
observed working a trap nest on June 17 and June 22. As of August 26,
106 trap nests showed nesting activity: Megachilids - 12: Eassaigecus
- i: eumenids. mun. Trypoxylonlni, W and Pompilids - 93.

Site II was located in 100 meters of forest edge, between a red
pine plantation and an old field. northeast of Pine Brae faculty

housing. Sixteen bundles of pine trap nests, prepared after a

 

 

108

technique described by Fye (1965) were placed at this site. Bore
diameters of trap nests were 3.2 an: and 4.8 mu. B. W Smith
females were observed working three days in June: one on June 20,
three on June 23. and three on June 29. As of August 26. 142 of 144
trap nests were in use: megachillds - 18: W - 27: eumenids,
Trypoxylonlni and others undetermined - 91.

Site III was a snail clearing adJacent to the Concordia ponds.
Nesting materials consisted of eimt 3 X 3 bundles of pine trap nests
with bores of 3.2 um and 4.8 m: and 72 Rims cuttings 150 um long. As
of August 26. 1983. 69 pine trap nests were in use: megachillds - 3:
Passalgegus - 12: eumenids, Psenini and Trypoxylonlni - 54. No
nesting activity was found in Rim: or Rum: cuttings.

Site IV was a south-facing slope overlooking the Huron River.
The slope is bordered on the east by a small pine plantation and on
the north by a mixed hardwoods. The slope is an old field covered
with grasses and annual weeds. Rhys is established along the
plantation edge and saplings of W, Juglans and mu, along
with mature W are scattered over the area. One hundred
forty-four pine trap nests with bore diameters ranging from 3.2 m to
8.0 m were distributed at this site along with thirty-six 150 ll!!!
Ruby: cuttings and thirty-six 150 um Engaging: twigs. One hundred
eight of 144 pine trap nests were used: megachillds - 11: We:
- 4: Win - 3: and eumenids, Psenini, and Trypoxylonlni - 90.
One Rum: cutting was excavated and eleven Emmy: twigs had mud

plugs.

Q.

 

109

Site V was in a 100 meters of fence row bordering the western edge
of the campus, south of Geddes Road. The plant cover at this site
consisted of Rhys, Rgbinia, and grasses. Nesting material included 54
pine trap nests with bore diamters of 3.2 mm to 8.0 mm, 108 Eraxinga
twigs, 27 Bugs twigs, and 27 plastic straws. As of August 26, 1983,
forty-five pine trap nests were used: megachillds - 2: Eassalgegus -
1: lsgdgntia - 9; others - 32. Twenty-five Eraxings were used and no

activity was observed in Rugs cuttings or plastic straws.

 

 

110

 

 

Huron River

Figure 16. Schematic map of Concordia College. Ann Arbor. Michigan.
indicating trap-nesting sites for i983.

111
Table 29 sunlnarizes the frequency of use of various nesting
materials. These studies clearly demonstrate an abundance of
trap-nesting wasps on Concordia’s campus. Artificial nesting
materials. l. e. trap nests and Enaxings twigs. were highly efficient
in attracting trap-nesting wasps. Rhys, Mg, and plastic straws

were very ineffective.

Table 29

 

Frequency of Nest Use by Nest Material Type.

 

Nest Type Number Available Number Used % Used

 

 

 

Pine

trap nests 558 369 66.13
£5?§§nu§ 144 36 25.00
Rim: 171 o o
Kuhn: 144 0 0
Plastic

straws 2? 0 0

All Traps 1044 405 38.79

 

112

Passalgegua was the presumed user of 45 pine trap nests based
upon direct observations of wasps at trap nests. closures consisting
of resin. or resin and impressed aphids. Examination of the contents
of these trap nests and reared wasps confirmed Passalgeggs use in 23
trap nests. Bight male and six female Passaigegus cuspldatus smith,
five main, and four ichneumonlds were reared. Trap nest use by
W was limited to bore diameters of 3.2 m and 4.8 m, and
all Eassalgegua trap nests, confirmed on the basis of reared

materials. were from Site 11.

APPENDIX 3

We of Michigan (Nymenoptera: Sphecidae)

Introduction

The genus Eassalggmma consists of small (4-9 mm) black wasps of
the Pempredeninae that provision their nests with aphids. The genus
is in the tribe Pemphredonlnl by virtue of the presence of two
submarginal cells in the forewings and antennal sockets placed near
the fronto-clypeal suture. The presence of three discoldal cells. two
recurrent veins and a relatively small stlgna place We in the
sub-tribe Pewhredonina. W may be distinguished from other
genera. of this sub-tribe by the following characteristics: inner
margins of compound eyes parallel or weakly converging below: scapal
basin weakly depressed: mandible with two or three weak teeth: ‘labrum
nearly triangular with a pronounced rounded apex: pronotum with a
strong transverse carlna: episternal sulcus well defined:
hyposternauius horizontal and the petiole shorter than broad (Bohart

and Nenke, 1976).

113

 

114

Recent taxonomic studies have increased the number of recognized
Nearctic species from seven to sixteen. Bohart and Menke (1976) list
seven Nearctic and four Holarctic species of Passalgegus. Muesebeck.
C.F.V., et. al. (1979) list eleven W in America north of
Mexico. Vincent (1978) recognized thirty-five species. sixteen as
North American, of which seven were known to occur in Michigan. Ne
did not include 2. mm Dahlbom though a single male has been
reported by Krombein (1961b) from Michigan. Bight species of
W are thus known to occur in Michigan. Specimens of
W were examined from the Museum of Zoology of the University

of Michigan and the Entomology Museum of Michigan State University.

Biology of North American Passaigecus

Passalgegus are known to nest in decayed wood. abandoned beetle
borings, stems of Rims, Rgsa, W, nga. bark of pine, cedar.
and oak. Nests are linear and usually partitioned and closed with
pine resin. All Passalgegns provision nests with aphids and a single
Passalgegua species may prey on more than six different aphid species
(Fye, 1965). Dixon (1973) states that aphid predators are more
closely associated with particular habitats than they are to
particular aphids. W could then be described as general
opportunists.

Passalgegus are markedly protandrous., Emergence occurs in late
May and June. Mean emergence for males is nearly one week earlier
than females. A few days after emergence females are observed
searching for nesting sites. Preferred sites are found in moderate
plant cover with nests in partial shade. Previously used nests are
cleaned of debris and reused. New excavation is limited to decayed
wood or pith. Generally, innermost cells produce females: outermost
cells produce males. Data vary on the relationships between the sex
of cell occupant. size of cell, and quantity of food stores. Eggs are
placed on the thoracic and abdaninal ventor of an aphid on the top of
the food mass. Larval feeding is completed within 40 days (Fye, i965)
and Baasalgeggs, over winter as prepupae. Pupatlon of Nearctic

materials occurs in late April and May.

115

 

Aphid Prey of W

Dixon (1973) notes that aphid parasitoids are fastidious, often
preying on a single species, but predators feed on a number of
species. Aphid prey of W are given by Fye (1965) and
Krombein (1955. 1956. 1958. 1960, i961a. 1967). W are
apparently generalists in their prey selection. The number of prey
provisioned by Eassalgegus is extremely varied. Prey provisioned per
cell ranged from seven to sixty-three: prey per nest ranged from fifty
to two hundred. Corbet and Backhouse (1975) suggest that in favorable
conditions a single female could kill fifteen hundred aphids based
upon one cell provisioned per day (thirty aphids) and a hunting life
of fifty days. No data have been given indicating mean nunbers of
aphids per cell. nor has any information been given on whether a
relationship exists between aphid species provisioned and mean number

of provisions.

116

 

Key to Michigan Passaigecus

Readers are referred to Vincent (1978) for a discussion of the
morphological features inportant in distinguishing We
species. A generalized diagram of the scutum and mesopleuron
illustrating structural features and terminology used in descriptions
and the following key may be found in Figs. 1 and 18. The following

key is an adaptation of Vincent’s Key to North American Raw
Shuckard to Michigan Eassalgegus.

1. Males: eleven flagellomeres: seven visible gastral

terga: clypeus with dense covering of silvery setae:

length of scape 2 times width ............. 2
1’. Females: ten flagellomeres: six visible gastral terga:

clypeus with sparse covering of silvery setae: length

of scape 3 - 4 times width . . . . . . . . . . . . . . 9

2. Spinose tubercles present medially on gastral tergum VI . . 3

2’. Spinose tubercles absent on gastral tergum VI . . ..... 7

3. Clypeal lobe tridentate (Fig. 8): apex of foretarsemere
produced below . . . . . . . . . . . . . . E. cuspldatus

3’. Clypeal lobe truncate or weakly concave .......... 4

4. Scrobal sulcus absent or weakly impressed (Figs. 19, 23) . . 5
4’. Scrobal sulcus deeply impressed. areoiate (Figs. 20 - 22). . 6

117

 

5’.

6’.

7’.

8’.

9"

118

Apical 1/3 to 1/2 of flagellmeres III - IX white
or pale yellow: hind trochanter pale amber . . B. W

Flagellomeres, trochanters dark brown

orbiack .......... .. KW

Mandibles. trochanters dark brown or black . . . . 2. mm:
Mandible white or pale yellow:
trochanters pale amber ........... E. W

Onaulus absent: scrobal sulcus absent: pronotal lobe
and trochanters dark brown or black . . . 2. mm
Onaulus present (Fig. 8): scrobal sulcus impressed,

areoiate . . . . . . . . . . . . . ........... B

Scutum anterioraliy with interrupted transverse carinae:

notaull longer then adnedian lines, extending to

mid-point of scutum (Fig. 2). . . . . . . . . . E. gracing
Scutum with course punctatlon, lateral margins with

weak transverse carinae: notaull not longer than

adnedian lines, terminating before mid-point of
scutum (Fig. 6) ................ 2. mm

unaulus present (Fig. 8) .................. 10
Onaulus absent ...... . ....... . . . ..... 11

 

10.

10’.

11.

11’.

12.

12’.

13.

13’.

14.

14’ O

119

Scutum anteriorly with interrupted transverse carinae (Fig. 2)
no setal division pattern on face: notaull extending to
mid-point of scutum: basal 1/4 of hind tibia creamy
white . . . . . ............... E, gnagilis

Scutumlwith course punctation and lateral margins with
weak transverse carinae: notaull not extending to

mid-point of scutum.(Fig. 6): tibia black . . . E, tunignum

Scrobal sulcus strongly impressed or areoiate ....... 12

Scrobal sulcus not present, or weakly impressed ...... 14

Scrobal sulcus strongly impressed: clypeal lobe
tridentate (Fig. 8): pronotal lobe and basal 1/3 to 1/4 of
hind tibia white ....... ’ ...... 2. We
Scrobal sulcus areoiate:

clypeal lobe truncate (Figs. 10, 13) .......... 13

Clypeal lobe width 1/4 of clypeal base width:

trochanters amber . . . . ..... . . . . . E. azeglatug
Clypeal lobe width one-fifth clypeal base width:

trochanters brown to black .......... E. lingatg:

Clypeal lobe weakly tridentate and weakly upturned (Fig. 12):
pronotal lobe. labrum and trochanters black. . P. singulanis

Clypeal lobe concave or convex .............. 15

 

15.

15’ .

120

Ciypeus concave (Fig. 7): pronotum white: labrum brown:

trochanters black . . . ......... . E. W

Ciypeus convex (Fig. 14): labrum and
trochanters amber .............. E. W

Michigan W

PASSALOECUS ANNULATUS (SAY)
(Figs. 14. 15, 23, 24)

PEMPHREDON ANNULATUS SAY, 1837. Boston Jour. Nat. Hist. 1:379.

REMARKS: Females: clypeal lobe proJecting strongly with margin weakly
convex: pronotal lobe and paipi white: labrum, mandible and legs pale
amber: males: apical 1/3 to 1/2 of flagellomeres I-X pale amber.

BIOLOGY: The biology of W W (Say) is reported by
Krombein (1955, 1958, 1960, 1961a, 1963) and Vincent (1978). Nesting
sites included anobiid borings, beetle borings in wooden cowshed
walls, twigs of Rugs glam L. and Man: um L. respectively, and
a boring in a red cedar stump. Data are given on four nests. One
pine trap nest contained three cells - 6.0, 10.0, and 12.0 m long
respectively. Two males were reared from this nest. One nest from
Eng: glabna L. and another from Juglans nigna L. collectively
contained six brood cells ranging from 10 mm to 30.7 mm long. Three
adults. one male and two females were reared. Three others died in

the prepupa stage. One nest had two vestibular cells, 19.0 mm and

121

 

122

16.0 nan long. Larval cocoons usually consisted of two transverse
silken partitions, the first contiguous with the resin partition and
the second 4.0 to 10.0 m distant from the first. Krombein (1963)
reported 2. annulatus active from May 20 to October 13 and considered
it multivoltine in the Washington, D.C. area. Vincent (1978) reported
that collection records indicate E. annulatgs is univoltine in
Missouri.

MICHIGAN DISTRIBUTION: Fig. 23

MICHIGAN FLIGHT DATES: 5 June - 16 Aug.

PARASITES: Unknown.

PREY RECORDS: Drepanaebis (Krombein. 1955. 1958): Macrgslpnum
(Krombein, 1958, 1960): Negtbgmasia (Krombein, 1961a).

123

PASSALOECUS AREOLATUS VINCENT
(Figs. 3, 13, 21, 25)

PASSALOECUS AREOLATUS VINCENT. 1978. Wassman Jour. of Bio. 36:159.
REMARKS: Females: clypeal lobe truncate: scrobal sulcus areoiate and
strongly impressed: notaull areoiate and slightly longer than adnedian
lines: all trochanters, tibia, and tarsi of foreleg amber: males with
slightly darker scape and mandibles. The light color of the
trochanters contrasts dramatically against dark brown femur.
BIOLOGY: Eaggglgggua yapgglgtgs Vincent was reported by Krombein
(1958, 1963) as E. relatiygs Fox. One female was observed carrying an
aphid on May 30, 1957 and June 1, 1957. Wasps were observed at beetle
borings from May 21 to June 2. Vincent (1978) reported on two nests
in pine trap nests with 1.5 mn borings from Bolivar, Polk County.
Missouri. Both had resin partitions and‘closures of resin and wood
particles. One nest had four cells with lengths of 8.3, 8.5, 9.0 and
8.5 mm and two vestibular cells: one male 2. aneglatgs and a chrysidid
(annulus sp.) were reared. Others died during rearing. A second nest
had five cells with lengths of 6.8, 7.0, 6.0, 15.5, and 16.0 mm. The
four outermost cells had been destroyed and a single female 2,
aneglatgs was reared from cell number one.
MICHIGAN DISTRIBUTION: Fig. 25
MICHIGAN FLIGHT DATES: 25 July - 9 Aug.
PARASITES: A chrysidid. annulus sp. (Vincent. 1978).
PREY RECORDS: Unknown.

124

PASSALAOECUS CUSPIDATUS SMITH
. (FIGS. 8, i9, 26)

PASSALOECUS CUSPIDATUS SMITH. 1856. Cat. Hym. Brit. Mus. v.4:427
RfliARKS: Females: clypeal lobe strongly tridentate: scrobal sulcus
impressed, with weak longitudinal carinae: pronotal lobe, ventral
surface of scape, basal 4/5 of mandible. labium, palpi, and basal 1/8
- 1/4 of hind femur white: males similar except tarsi and pre-tarsi
of fore- and midleg amber. E. W is the largest of the
Michigan Easaaigegus.

BIOLOGY: Records of Passalgeggs ggspigatns Smith are provided by Fye
(1965), Krombein (1956, 1958, i963. 1967), and Vincent (1978).
Nesting sites included beetle borings in wooden cowshed walls,
artificial borings in elderberry and chinaberry, pine trap nests, and
soda straws. Krombein observed nesting activity near Arlington,
Virginia May 23 - May 29. 1956: May 30. 1958: May 17 - May 30, l959:
June 1 - June 7. l960: May 22 - June 1. l962: and May 19, l963.
Nesting activity was observed near Derby, New York June 7 - July 4,
1959 and June 1 - July 9. 1960. Twenty nests were in pine trap nests
with a bore diameter of 3.2 mm. These nests contained 58 provisioned
cells. Cell lengths were 8 - 52 um (mean = 16.3). Eighteen female
cells were 12 - 47 mm long (mean = 16.1) and 14 male cells were 10 -
31 m long (mean = 14.8). Two nests from 4.8 m borings had 4
provisioned cells respectively 7, 8. 13. and 126 mm long. One 6.4 mm

boring had 4 provisioned cells 7, 6, 7, and 9 m long. No intercalary

 

125

cells were found and all but one nest had a vestibular cell.
Vestibular cells were 5 - 145 um long. Resin partitions were usually
.25 um thick with one occasionally 4 m thick. Resin closures were
usually 1 nm thick and ranged from .25 to 4 um. Krombein (1967)
reported that E. W had a larval feeding period of 6 to 9
days. Fye (1965) found 2. We in two 6.4 nm borings in
eiderberry or chinaberry. One nest contained 4 cells: females were
reared from 3 cells (11 - 21 m. mean = 15: and one cell (11.2 m)
produced a male. The vestibular cell (102 m) was closed with a 3.2
an resin plug. A second nest of five cells (6.4 - 14.4 illil) prodiced
females from one 6.4 m and one 8.0 m cell. The vestibular cell was
30 ll!!! long, partitioned and closed with a 1.6 m resin plug.

Vincent (1978) observed two females cleaning debris from old nests
and provisioning nests August 12, l972 at Willard’s Peak, Utah
(9500’). Prior to provisioning. wasps ringed nest openings with two

drops of resin. Eight provisioning trips were observed. Trip

duration was 3.8 - 10 minutes (mean 5.96 1 2.85) Thirteen resin
gathering trips were observed with durations of 0.5 - 8.0 minutes
(mean = 2.04 1 1.93). Nest closure was achieved with four drops of
resin spread diagonally across the nest opening. Nest architecture
was reported from 83 soda straw nests from Montclair. Alameda Co.,
California. Straws measured 90 an x 4 Hill and contained two to nine
cells (mean = 6.14 1 1.75). Five hundred-ten brood cells were
examined. One hundred-eleven produced females. Cells were 5.0 - 18.0

um long (mean = 10.09 _+_ 2.19). One hundred-ten male cells were 4.0 -

16.0 run long (mean = 8.82 1; 2.16 m). Indivimal cells contained 7 —

“a *1

126

32 aphids. Resin partitions were 0.5 - 4.0 nm. Vestibular cells
werefound in all nests (3.0 - 21.0 um long, mean = 9.89 .+_ 3.5, n =
108). Flfty- five nests had one vestibular cell, twenty-two nests had
two, and three cells had three. Resin closures had coverings of
coarse wood particles.

MICHIGAN DISTRIBUTION: Fig. 26.

MICHIGAN FLIGHT DATES: 13 June - 14 Aug.

PARASITES: Two chrysidids: mm: m (Fabricus), (Krombeln, 1967:
Vincent, 1978) and W W (Vincent, 1978): two
ichneumonlds: Rgmenia W (Cresson), (Krombein. 1967): and
Romania 3mm (Cresson), (Vincent. 1978)

PREY RECORDS: mu, (Fye, 1965): W, (Fye, 1965: Kramein.
1956, 1958, 1967): Mugs (Kraubein, 1967): M32013. (Kronbein,
1967): 81152931911211.0111. (Fye. 1965).

127

PASSALAOECUS GRACI LI S ( CURTI 8)
(Figs. 2. 11. 22)

DIODONTUS GRACILIS CURTIS. 1834. British Entomology 11:496
REMARKS: Females: clypeal lobe concave, scrobal sulcus areoiate:
scutum with oval raised scutal patches: notaull areoiate. extending to
midpoint of scutum: scutum anteriorly with transverse interrupted
carinae: areoiate omaulus narrowly separated from episternal sulcus:
mandible. except apex. ventral surface of scape, pronotal lobe.
foretibla and basal 1/4 of mid— and hind tibia creamy white. This
species is very uncannon and the transverse carinae of the scutum are
distinctive.
BIOLOGY: Unknown.
MICHIGAN DISTRIBUTION: One female: Wexford Co., 20 August 1973.
collected by R. D. Averill: deposited in the Entomology Museum of
Mighigan State University. Two males: Macomb Co., 19 March 1961,
collected by S. J. Thomas, deposited in the USNM.
MICHIGAN FLIGHT DATES: 20 Aug.
PARASITES: Unknown.
PREY RECORDS: W, (Danks. 1971): W, (Danks. 1971):
Mahala, (Danks. 1971).

128

PASSALOECUS LINEATUS VINCENT
. (Figs. 5, 10, 27)

PASSALOECUS LINEATUS VINCENT, 1978. Wassman Jour. of B10. 36:162
REMARKS: Females: clypeal lobe narrow and truncate: scrobal sulcus
areoiate and strongly impressed: notaull areoiate extending to
midpoint of scutum: pronotal lobe. ventral surface of scape, basal 2/3
of mandibles, and palpl white: trochanters dark brown: males similar.
BIOLOGY: Unknown
MICHIGAN DISTRIBUTION: Fig. 28.
MICHIGAN FLIGHT DATES: 13 June - 20 July
PARASITES: Unknown.
PREY RECORDS: Unknown.

129

PASSALOELCUS MONILICORNIS DAHLBOM
(Figs. 7, 28)

PASSALOECUS MONILICORNIS DAHLBOM, 1842. Dispositio methodica:12
REMARKS: Females: clypeal lobe weakly concave or truncate: scrobal
sulcus 'weakly impressed or absent: notaull areoiate. as long as
adnedian lines: pronotal lobe. ventral surface of scape, basal 4/5 of
mandile white: trochanters and femurs dark brown: tarsi and pre-tarsi
light brown: males similar.

BIOLOGY: Fye (1965) and Krombein (1967) described nesting habits of
E. Wm}; Dahlbom. Fye reported on thirty-seven nests in
drilled eiderberry twigs from the Black Sturgeon Lake area of
northwestern Ontario. Nest diameters were 6.4 II!!! and 8.0 am with 6.4
mm preferred. Female cells were 4.8 - 12.8 mm long (mean = 8.66 1.5
um), male cells were 4.8 - 11.2 run long (mean = 7.8 1.3 um). Five
nests of 6.4 mm boring had 5 - 10 cells each (mean = 7). Vestibular
cells were 1 - 4 per nest. 26 - 77 mm long. Partitions and closures
were of resin and closure plugs were 1.6 - 3.2 mm thick. In 8.0 mm
borings mean length of female cells was 10.0 m. A single male cell
was 14.4 mm long. Unlvoltine and bivoltine populations were reported.
The univoltine population provisioned nests in late June. The larval
feeding period lasted up to 40 days. Pupatlon occurred the first and
second week of the following June with emergence the last week in
June. The bivoltine population provisioned nests in June: larval

feeding was completed by mid-July and adults emerged July 29 - August

' 55.

130

4. Nests provisioned by this summer generation were collected from.16
August - 3 September and adults of the overwintering generation
emerged the last week in June.

Krombein (1967) reported on six E.mgn111ggnni§ nests in pine trap
nests near Derby. New York. Four nests with 3.2 mm borings contained
20 provisioned cells 6 - 12 mm long (x = 8.6mm). Twelve female cells
had a mean length of 9.6 mm and four male cells had a mean length of
6.7 um. Three of these nests contained six cells each and a fourth
nest contained two cells. A nest with a 4.8 run bore contained 12
stored cells 6 - 12 run long (x = 8.6) and a second 4.8 m nest had
three stored cells 19, 25, and 90 mm long respectively. No nests had
intercalary cells and vestibular cells were 10 - 20 am long. Resin
partitions were .25 mm - .50 mm and closures were 0.5 - 1.0 mm thick.
Kroubeln considered 2. W bivoltine in this area. Adults,
from four nests completed in late May and early June, emerged June 23
- July 18. A single adult emerged August 17 from a nest stored in
mid-July. One nest, containing six completed cells, produced females
from three inner-most cells, cells four and five produced males and

cell six suffered egg death.

131

MICHIGAN DISTRIBUTION: Fig. 28.

MICHIGAN FLIGHT DATES: 24 May - 8 Aug.

PARASITES: Two chrysidids: mm; m (Fabricus), (Fye, 1965) and
annulus 131mg ? (Norton), (Fye, 1965): two ichneumonlds: mm
mm (Cresson), (Fye, 1965) and mm; W ? (Cresson),
(Fye. 1965).

PREY RECORDS: mum, (Fye, 1965): Anuraphis, (Krombein. 1967):
mm. (Fye, 1965): W. (Fye, 1965): Will. (Fye,
1965): machine. (Fye, 1965): Elem ?. (Fye, 1965):
Wm (Krombein, 1967)

132

PASSALOECUS SINGULARIS DAHLBOM
. (Figs. 4. 12. 16, 29)

PASSALOECUS SINGULARIS DAHLBOM. 1844. Hym. Europaea, v.1: 243
REMARKS: Females: clypeal lobe weakly upturned and weakly
trl-dentate: scutal patches present: no white markings: males similar.
This species is the slenderest of the Michigan Passalgggga.
BIOLOGY: Unknown.
MICHIGAN DISTRIBUTION: Fig. 29.
MICHIGAN FLIGHT DATES: 17 May - 30 Aug.
PARASITES: Unknown.
PREY RECORDS: Unknown.

133

PASSALOECUS TURIONUM DAHLBOM
(Figs. 6. 9. 17. 20)
DIODONTUS GRACILIS CURTIS. 1834. British Entomology 11:496
REMARKS: Females: clypeal lobe concave: scrobal sulcus strongly
areoiate: omaulus present: notaull areoiate. as long as adnedian
lines: scutal patches present: ventral surface of scape and basal 2/3
of mandible dark brwon to pale white. This species like 2. gzagillia
possesses an omaulus. but lacks the transverse carinae of the scutum.
BIOLOGY: While conducting preliminary research on the trap-nesting
biology of Eaasalgegga five unusual females (one - 6 August 1982, one
- 27 July 1983, two - 1 October 1983. and one - 30 August 1985) were
collected. Using Vincent’s revision of the We of North
America (Vincent. 1978) this material was identified as W
0923.11.11 Dahlbom. Subsequent to the use of Vincent’s key this
material was cmpared with a description of 2. 1291291111 and a specimen
previously determined as 2. mm from the Michigan State
University Entomology Museum Collection. The determination of this
material was placed in doubt because of deviations from Vincent’s
description and distribution records for m that showed the
eastern limit of its range to be Colorado. During a subsequent review
of Yarrow’s paper on nanenclatorial problems among European species of
Eassalgegu: the present material was clearly described and identified
as W m Dahlbom (Yarrow, 1970).
Kroubein (1961) reported on two male W reared from a
twig from Macoub Co. Michigan. Althoud: positive identification was

not possible he identified the material as standing close to

134

B. We Morawitz. A specimen was sent to Dr. J. deBeaumont,
Lausane, Switzerland. He identified the material as E. W
Dahlbom for which he considered 2. W a synonym. Kroubein
considered 2. tgnlgngm.0ahlbom adventive in the United States since no
records existed for this material prior to July 1941. Vincent (1978)
assigned this material to B. gnagillxs (Curtis).

Yarrow (1970) untangled the nomenclatorial problems among names
applied to We spp. by British and Continental authors, and
the confusion of We types. He clearly distinguished three
species of We that possess an areoiate omaulus anterior to
the episternal sulcus: E. gracing (Curtis). E. magnum Dahlbom, and
E. banenlla Dahlbomn

Vincent (1978) agreed with Yarrow’s interpretation of this group
and raised 2. magnum Dahlbom from synonOmy with E. mun (Curtis)
to species status. However, he did not provide a description of this
adventive species.

DESCRIPTION: Female. black: ventral surface of scape white. basal
half of the anterior face of mandible light amber: pronotal lobe
amber: fore-tibia above light brown: lablum, posterior face and distal
half of mandible. pro-. meso-, and meta-tibia. pro-. meso-. and
meta-femur dark brown: setal division pattern low on face at level of
antennal sockets: clypeal setae on dorsal 1/2 inclined toward midline,
on ventral 1/2 inclined ventrally: clypeal lobe weakly concave:
interocellar distance equal to ocellocular distance: notaull impressed
and areoiate. as long as adnedian lines: scrobal sulcus areoiate:
scutal margin opposite tegula strongly reflexed and areoiate: scutum

with dense, course punctation: posterior margin of scutum areoiate

 

135

with inner margins of fovaea open: lateral faces of pronotum with
strong longitudinal carinae: dorsal surface of pronotum posterior of
pro-notal carina areoiate: omaulus present, nearly contiguous with
episternal sulcus.

This material is differentiated from 2. bgmLLa Dahlbom by the
following characters: head height/head width = 1: lack of white or
amber markings on legs: dense. coarse punctation of the scutum: strong
areoiation of lateral margins of scutum (Fig. 6): distinct lateral
carinae and areoiate dorsal surface of the pronotum. Hing characters
also differ from 2. mm. The distal portion of the marginal cell
is narrowly pointed, the first radial-medial cross-vein is distinctly
bent mid-way between the radius and medius: and the basal vein is
strongly arched (Fig. 17). The gaster is differentiated by the
angulate posterolateral margin of the gastral tergum II.

The material under consideration is placed in Vincent’s 'Gracilis
Group - Group Assemblage II' on the basis of the following characters:
females with scutal patches: concave clypeal lobe: and well-developed
omaulus and scrobal sulcus (Fig. 20).

BIOLOGY: Unknown.

MICHIGAN DISTRIBUTIGV: Five females. Hashtenaw Co., one each August
6, 1982, July 27, 1983, August 30, 1985, and two October 1, 1983:
deposited in the Entomology Museum of Michigan State University. One
female. Marquette Co., 23 June: deposited in the Museum of Zoology of
the University of Michigan

MICHIGAN FLIGHT DATES: 27 July - 1 Oct.

PARASITES: Unknown.

PREY RECORDS: Unknown

Figure

Figure
Figure
Figure
Figure
Figure
Figure
Figure
Figure
Figure
Figure
Figure
Figure
Figure
Figure
Figure
Figure

Figure

Figure
Figure
Figure
Figure

Figure

10.
11.
12.
13.
14.
15.
16.
17.
18.

22.
23.

136
LIST OF FIGURES (APPENDIX 3)

Dorsal view of the Scutum Illustrating structural
features and the terminology used in descriptions

and keys (after Vincent. 1978) ............. 138
Dorsal view of the scutum of E. gmagilia ....... 138
Dorsal view of the scutum of E, angglatgs ....... 138
Dorsal view of the scutum of E. singulanis ...... 138
Dorsal view of the scutum of P, Lineatus, ....... 138
Dorsal view of the scutum of 2, tynlgngm ....... 138
Clypeal outline of E. maniliggnnis .......... 139
Clypeal outline of P. guspigatgs ........... 139
Clypeal outline of 2, tunigngm, ........... 139
Clypeal outline of E. lineatus ........... 139
Clypeal outline of Rt gnagilis ........... 139
Clypeal outline of E, singulanis ........... 139
Clypeal outline of E. areoiatus ........... 139
Clypeal outline of E, annulatga ........... 139
Forewing of 2, annulatgs .............. 139
Forewing of 2. singulanis .............. 139
Forewing of 2. 1.11m .............. 139

Lateral view of left mesopleuron illustrating
features and terminology used in descriptions

and keys (after Vincent, 1978). ........... 140
Left mesopleuron of P, ggspiganua .......... 140
Left mesopleuron of 2. tunignum ........... 140
Left mesopleuron of E, areglamma ........... 140
Left mesopleuron of 2. gnagilis ........... 140
Left mesopleuron of P. annulatus .......... 140

 

Figure 24.
Figure 25.
Figure 26.
Figure 27.
Figure 28.

Figure 29.

Michigan
Michigan
Michigan
Michigan
Michigan

Michigan

distribution
distribution
distribution
distribution
distribution

distribution

137

of E. annulatus ........ 141
of 2, angglafiua ........ 141
of E, nggldagua ........ 141
of 2. uneasy: ........ 141
of E. mgniliggnnis ....... 141

138

Admedian Line
\ ’Notaulus

   

Parapsidal Line

Scutal Patches

 

P. singularis P. lineatus

 

P. turionum

 

l

7

P. cuspldatus

W

P. lineatus

P. monilicornis

10

   

P. turionum

 

 

mm

P. QVQCIIAS P. sinaularis
13 ”mm 14 $
P- areoiatus P. annulatus

 

 

 

    

 

.. \VV /
16
P. singularis

 

 

P. turionurn

um
Scrobal Sulcus

Episternai

 

Hypersternalus

N p—el

 

 

 

P. cuspldatus P. turionum

 

P. areoiatus P- annulatua

141

 

 

 

 

 

P. monilicornis P. singularis

142

Literature Cited

Bohart, R. M.. and A. S. Menke. 1976. Sphecid wasps of the
world. ix + 695 pp. University of California Press.

Corbet. S. A.. and M. Backhouse. 1975. Aphid Hunting Wasps:
A Field Study of Passalgeggs. Tran. Roy. Ent. Soc.
London. 127: 11-30.

Dixon, A. F. G. 1973. Biology of Aphids. The Institute of
Biology’s Studies in Biology no. 44, 58 pp. Edward Arnold
(Publishers) Limited, London.

Fye, R. E. 1965. The Biology of the Vespidae, Pompilldae and
Sphecidae from trapnests in northwestern Ontario.
Canadian Ent. 97: 716-744.

Krombein. K. V. 1955. Miscellaneous prey records of solitary
wasps. 1. Bull. Brooklyn Ent. Soc. 50: 13-17.

. 1956. Miscellaneous prey records of solitary
wasps. II. Bull. Brooklyn Ent. Soc. 51: 42-44.

 

1958. Miscellaneous prey records of solitary wasps.
III. Proc. Biol. Soc. Washington. 71: 21-26.

1960. Biological notes on some Hymenoptera that nest
in sumach pith. Ent. News. 71: 29-36. 63-69.

1961a. Miscellaneous prey records of solitary wasps.
IV. Bull. Brooklyn Ent. Soc. 56: 62-65.

1961b. Eassaimus .turipnum Dahlbom. an

adventive European wasp in the United States. Ent.
News. 72: 258-259.

 

‘_____. 1963. Natural history of Plummets Island Maryland.
XVII. Annotated list of the wasps. Pro. Biol. Soc.
Washington. 76: 255-280.

. 1967. Trap-nesting wasps and bees: life histories.
nests, and associates. iii-vi + 570 pp. Smithsonian
Press. Washington D.C.

 

Muesebeck. C. F. W.. K. V. Krombein. et. al. 1979.
Hymenoptera of America north of Mexico: synoptic
catalog. Smithsonian Institution, Washington, D. C.

143

Vincent. 0. L. 1978. A revision of the genus Passalgegua
(Hymenoptera: Sphecidae) in American north of Mexico.
Wassmann J. Biol.. 36(1-2):127-i98.

Vincent, D. and J. D. Hoffman. 1974. Advantages of using
fluorescent light with dissecting microscopes in
taxonomic investigations. Ann. Entomolo. Soc. Am.,
67:820-821.

Yarrow, I. H. H. 1970. Some nomenclatorial problems in the
genus Passaigegus Shuckard and two species not before
recognized as British. Ent. Gazette, 21:167-189.

 

 

APPENDIX 4

Data Tables from Trap-nesting Studies of Eassalgeggs. 1984-1987

 

Table 30.

 

Rearing data for We cuspldatus. 1984-

 

 

 

 

 

 

Bore Parasitized Otherwise Adults
(mm) Cells cells failed cells Prepupae Reared
3.2 78 10 5 63 30
4.0 29 7 4 18 9
4.8 52 8 4 40 31
6.4 10 3 i 6 5
Totals 169 28 14 127 75

 

 

144

145

Table 31.

 

Bore diameters and lengths of Eassalgeggs ggspidatus cells, 1984.

 

MEAN
MEDIAN
S. D.
S.E.M.

MEAN (A)

MEDIAN (A)
S. D. (A)
S.E.M. (A)

Bore Diameter (mm)

 

3.2

13.5443
12.07
4.7702
.5366?

N = 79

12.5811
11.90

3.0359

.3529

N=74

4.0

19.5862

11.125

23.4028
4.3458

N = 29

11.5652
10.00

2.3527

.4906

N = 23

4.8

17.1154
9.6428

23.853
3.3079

N = 52

9.4783

9.375

3.5737
.5269

N = 46

6.4

7.30

5.9286
4.5837
1.4495

7.30

5.9286
4.5837
1.4495

N = 10

 

(A):

Data were adJusted by a deletion of 10% of cell length values
from skewed end of distribution for pooled sample of all bore

diameters.

 

 

Table 32.

 

Frequency of trap nest use and number of adults reared out
for four We spp.. 1985.

 

 

 

 

 

 

Bore (mm)
1.6 2.4 3.2 4.0 4.8
We spp.
E. annuiatus (Say) 2(2) 3(6) -- -- --
E. ggsgigatgs Smith -- -- 2(5) 2(5) 3(12)
2. W Dahlbom 1(1) 5(13) -- -- --
E. simian: Dahlbom -- 1(3) —- -- --
Table 33.
Frequencies of trap nest use by four Paasglgegus spp., i986.
Bore dlaeters (I)
1 6 24) :L4 21! 2L2 315 (L0 4.4 1L8

Spence ___ ___ ___ ___ ___ ___ ‘___ ___ ___

:umuuuua i i 4 0 0 0 0 0

mining is 20 9 o o o o 0

My: 0 1 4 7 7 5 3 o

lanU30u05 i 3 8 2 2 0 0 (l

 

147

Table 34.

 

Basal cell lengths frm eimt bore dialeter classes and two bore depth
classes. Data compiled from four Eagsalgegga spp., 1986.

 

 

 

 

 

Bore depths (uni)
Diameter class
120 60 totals
Bore
diaeter Oil 8 Mean 01- 8 Mean 0:- 4 Mean
(I) BCLI BCH BCL BCL BC BCL BCL BC BCL
1.6 418 11 38 194 15 12.93 612 26 23.54
2.0 427 18 23.72 0 17 0 427 35 12.20
2.4 495 14 25.26 69 15 4.6 564 29 19.45
2.8 107 6 17.83 0 3 0 107 9 11.89
3.2 so 4 7.5 0 6 0 3) 10 3.00
3.6 0 6 0 0 6 0 0 12 0.00
4.0 25 5 5 0 2 0 25 7 3.57
4.4 0 2 0 0 i 0 0 3 0.00
Totals 1502 66 22.76 263 65 4.05 1765 131 13.47

 

l Basal Cell Length
H Basal Cell

 

148

Table 35.

 

Eaaaalgegu§.areglatus provisioned cell lengths. 1986.

 

(mm)

Bore diameter

 

len th
(mg)

Cell

2.4

2.0

1.6

 

 

_112212342112 1. 11 2
_ 1.4. 2 31135342125 1. 1.21. .12 2 2 1. I 1.
a... 1
11a 21. 2 al. 1

_ 2 1211.1. 733 22323 1.1

1 1111

nU.5nU.$050505050.50.50.505050505nw50505050sosososoososooooooooo

al.?”8899001122334HL556677889OH

11111111111111111111

m

OOOOOOOOOOOOOOOOOO

m112233445566778990
2222222222222222223

waxwg

 

149

Table 36.

 

Passaigegus cuspldatus provisioned cell lengths. 1984-1987.

 

Bore diameter (mm)

4.0 4.4 4.8 5.6 6.4

3.6

3.2

2.8

2.4

 

2.0

Cell
length
(mm)

— 2 7 11
9.
3 5 9 3161.0 8171.513 1 2
11 .1 1.. 1. 1.
2 2468m8fl4m4mlfl27 52722123 1
11311221112 1..

1. 5285231444351143810153514.1.3
2 2 1 2 1 1 1

3 414.13 322 3 1. 1.
2 3 31.71.7231. 1.11 2
1 1 1 1

5566778899001122334455667788990

111111111111111111112

 

 

 

150

Table 36., (cont’d)

 

Bore diameter (mm)

 

Cell

length

6.4

3.2 3.6 4.0 4.4 4.8 5.6

2.8

2.4

2.0

 

1111

ll . H|=I5
[

1

11 2 1 11 1 1 11111
1
1 1

3 1.. 2221.3 21 2 1. 1.

11 1 1 1

50505000000000000000000000000000000000000

.........................................

mam amufixmmmmmaxm

2.4.67
4.4.4.4.

1.5686
00001
11111

 

151

Table 37.

W W15 provisioned cell lengths, 1985-1986.

 

 

Bore diameter (Inn)

 

Ce] 1
length
(m)

3.6

3.2

2.8

2.4

2.0

1.6

 

 

‘.

4.1.1. 1.
1.3 21.

121.1. 21.
32415712224133.1141 1.

0505050505050s050505050505050500000000000000

OOOOOOOOOOOOOOOOOOOOOOOOOOOOOOOOOOOOOOOOOOO

556677889900112233445566778899 1234.5 7&9 228

11111111111111111111 22222 490
1.

 

15%!

Table 38.

 

Lengths of Eaagglge§¥s rovlsloned cells from ei t bore dlaleter classes
p Data were combined ggon four Ea§§a19g§n§,spp.,

 

 

 

 

 

Bore the (nu)
dep Diameter class
120 60 totals
e
diameter Cu- 8 of liean 0- t of llean Q- 8 of ilean
uII) I! Cells CL CL Cells CL CL Cells CL
1.6 397 26 15.27 456 29 15.72 853 55 15.51
2.0 768 48 16.00 570 46 12.39 1338 94 14.23
2.4 629.3 48 13.11 517 41 12.61 1146 88 13.02
2.8 338.5 30 11.28 75.5 8 9.44 414 38 10.89
3.2 142 13 10.92 102.5 ii 9.32 244 24 10.17
3.6 100 ii 9.09 113.5 14 8.11 213.5 25 8.54
4.0 172 16 10.75 23 2 11.5 195 18 10.83
4.4 59 5 11.8 14.5 2 7.25 73.5 7 10.50
Totals 2605.8 197 13.23 1872 152 12.32 4477.8 349 12.83

 

! Blghtf instances cells of extraordinary length were not included.
Daa frat hdeese letlons are snarized belaa.
l! Provisioned Cell Lengths

Bore depths (ml)

 

 

 

120 60

Bore
dialeter cu- I of lean Cue I of Hean
(ml) CL Cells CL CL Cells CL
1.6 126 2 63 80 2 40
2.0 100 i 100 -- -- --

2.4 -- —- -- 75 2 37 5

2.8 -- -- -- -- -- --
3.2 94 i 94 -- -- --
3.0 446 5 89.2 60 1 60
424 95 1 9s -- -- --

 

 

IEEB

Table 39.

 

Provlsloned cell lengths for four Eaaaalgegus spp., 1986. Cell
lengths are compared on the basls of bore dlaneters and bore depths.

 

llean cell lengths (In)

 

 

 

 

 

 

 

 

Bore depths (ll)
Bore
dlaleter
(II) All T’nests 120 60
Raggalgegus Hean (N) Hean (8) Mean (R)
app. ___... ___.
mum
All 11.7272 (22) 11.54 (11) 11.909 (11)
1.6 12.33 (3) - -~ 12.33 (3)
2.0 12.20 (5) -- -- 12.20 (5)
2.4 11.43 (14) 11.54 (11) 11.00 (3)
mm:
All 11.658 (120) 17.657 (73) 15.106 (47)
1.6 21.039 (38) 22.435 (23) 18.9 (15)
2.0 14.60? (61) 16.086 (35) 12.615 (26)
2.4 14.69 (21) 14 (15) 16.417 (6)
cuspldatus
All 14.287 (106) 16.463 (68) 10.1969 (38)
2.0 13 (4) 12 (1) 13.333 (3)
2.4 13.1 (10) 11.5 (2) 13.5 (8)
2.8 11.2857 (28) 11.8181 (22) 9.3333 (6)
3.2 9.1538 (13) 9.1 (5) 9.188 (8)
3 6 19.14 (21) 31.3 (10) 8.68 (11)
4.0 14.1818 (22) 14.1818 (22) -- ~-
4.4 21.0625 (8) 25.67 (6) 7.25 (2)
mammals
All 15.78 (50) 18.0776 (26) 13.2925 (24)
1.6 23 (2) -- -- 23.00 (2)
2.0 37.625 (4) 46.00 (3) 12.5 (1)
2 4 14.00 (22) 15.42 (6) 13.47 (16)
2.8 9.8125 (8) 9.8125 (8) -- --
3.2 9.9285 (7) 10.25 (4) 9.5 (3)
3.6 19.5 (7) 24 (5) 8.25 (2)

 

 

1E54

Table 40.

 

Provisioned cell len ths for four spp., 1986 with the
application of the 1 % exclusion ru e.

 

llean cell lengths (in)

 

 

 

 

 

 

 

 

 

 

Bore
Bore depths (mm)
diameter
(II) All t’nests 120 60
Eassg%gg§u§ Hean (N) Mean (N) lean (8)
minus
All 10.8 (20) 10.50 (10) 9.2 (10)
1.6 8.5 (2) -- -- 8.5 (2)
2.0 12.2 (5) -- -- 12.20 (5)
2.4 10.62 (13) 10.50 (10) 11.00 (3)
mm
All 13.94 (108) 14.134 (64) 13.4986 (44)
1.6 15.37 (31) 15.28 (18) 15.65 (13)
2.0 13.93 (59) 14.44 (33) 12.615 (26)
2.4 11.97 (18) 11.77 (13) 12.5 (5)
My:
All 10.13 (96) 10.7038 (59) 9.4189 (37)
2.0 13.00 (4) 12.00 (1) 13.333 (3)
2.4 10.22 (9) 11.50 (2) 9.86 (7)
2.8 11.23 (26) 11.3 (20) 9.333 (6)
3.2 9.1538 (3) 9.1 (5) 9.188 (8)
3.6 8.94 (18) 10.29 (7) 8.68 (11)
4.0 10.55 (20) 10.55 (20) -— --
4.4 9.42 (6) 10.5 (4) 7.25 (2)
mm:
All 11.72 (45) 10.3966 (24) 11.759 (21)
1.6 -- -- -- -- -- ~-
2.0 14.17 (3) 15 (2) 12.5 (1)
2.4 13.43 (21) 15.42 (6) 12.63 (15)
2.8 9.8125 (8) 9.8125 (8) -- --
3.2 9.9285 (7) 10.25 (4) 9.5 (3)
3.6 7.4166 (6) 7.000 (4) 8.25 (2)

 

 

155

 

 

 

 

 

 

 

 

 

 

 

 

Table 41.
“(Her of provisioned cells (rm eidit bore dineter classes
aggstwo bore c asses. Catined data frat four W spp.,
Bore ths (m)
dep llineter class
120 60 totals

Bore
diameter

(a) 8 Cl) T’NSTS1 liean 8 CI! T’ilS'i'S Bean 8 Cl! T’llSTS liean

1.6 28 11 2.55 31 15 2.07 59 26 2.27

2.0 49 18 2.72 46 1 2.71 95 35 2 71

2.4 47 14 3.36 43 15 2.87 90 29 3.10

2.8 m 6 5.00 8 3 2.67 38 9 4.22

3.2 14 4 3.50 11 6 1.83 25 10 2.50

3.6 16 6 2.67 14 6 2.33 30 12 2.50

4.0 16 5 3.20 2 2 1.00 18 7 2.57

4.4 6 2 3 00 2 1 2.00 8 3 2.67
Totals 206 66 3.12 157 65 2.42 363 131 2.77
111-oer of Cells/Mr of Trap llests

Table 42.

 

Frequencies of trap nest helmt selection by four W sup” 1986.

 

 

Heigit (II)
5 1 1.5 2
annuiatus 1 2 2 1
mlatua - 2 1 4
91121114111: 6 a 7 11
mum: 3 8 5 2

 

 

156

Table 43.

 

Vestibular cell lengths (VCL) frcl elgit bore diameter classes and two bore
depth classes. Data capiled (rm four W spp.. 1986.

 

 

 

 

 

Bore ths(-)
d” Diaeter class
120 60 totals

Bore
diaeterCl- ! liean I liean Cl- ! liean
(I) VCL VC VCL VOL 170 VCL VCL VC VCL
1.6 319.5 11 29.05 211 15 14.07 5&5 26 20.40
2.0 705 18 39.17 384 17 22.59 1089 35 31.11
2.4 371 4 26.50 213 15 14.2 584 29 an.“
2.8 251 6 418) 72 3 24 323 9 35.89
3.2 185 4 46.25 258 6 43 443 10 44.30
3.6 125 6 20.83 164 6 27.33 289 12 24.08
4.0 297 5 59.40 97 2 48.5 394 7 56.29
4.4 72 2 37.00 45 1 45 117 3 39.00
Totals 2327.5 66 35.27 1444 65 22.22 3771.5 131 28.79

 

 

157

Table 44.

 

 

 

 

 

Cell Data on Aphids Provisioned by Eassalgggg§_gu§21datu§, 1987.
Aphlds per Cell (cell 1 inner-most)

Aphid Trapnest
app.* number 1 2 3 4 5 6 7 8 9 10

1 1-85-116 32 38 25

1 2-86-18 53

1 1A-86-50 44

1 2A-86—24 36 37

1 3-86-85 41 41 26

1 2—85-096 30 27 26

1 2-85-083 31 43

1 4-85-049 36 52 40

1 3-6A-86-15 30 30 23 23

1 13-86-14 23 20 21

1 5-85-115 49 46

1 28-86-66 37 22 22 30

1 2-5A-86-62 31 44 29 30 30 28

1 3—6A-86-46 48 44 43 51 52

1 2-58-86-44 29 26 31 32 29

1 6—78-86—15 46 41

1 3—68-86-67 49 24 31 27 36

1 2-58-86—81 44 28 29 36 30 25

1 28-86-12 66 63 39 33

1 3-68-86-14 10 12 16 22 17 24 27 23 26 25

1 38-86-58 29

1 2-58-86—68 29 26 31 33 36 30 37

1 2-85-055 58 49 30 37 25

1 38-86-79 61 43 64 38 29 39
*1=111zzusmnandae:=2 CLnaniasmza=liacmsi2bum

21121192111333 4=

W89.andmsp:5=m
genus 6=mbb1ummnaez7=

W-

 

Table 44.. (cont’d.)

158

 

Aphld Trapnest

 

Aphids per Cell (cell 1 inner-most)

 

 

spp.4 number 1 2 3 4 5 6 7 8 9 10

1 2—5A-86-67 51 55 29 37 33

1 25-86-44 23 28

1 4n-86-31 44 38 26

1 28-86-32 51 27

1 38-86-21 36 so 30

1 3a-86-47 57 67 73 39

1 25-86-27 42

1 35-86-85 38 31 24 30 29 35

1 3-614-86-66 74

1 2-5a-86-07 26 26 35

1 2-5A-86-72 4o 49

1 3-68-86-20 9 29 14 17 25 35 36

1 28-86-82 34 41 ‘

1 3A-86-32 35 49 52 61 43 4o 46

1 38-86-60 47 41 53

1 3-68-86-07 47 53

1 01-86-68 47 53

1 3-68-86-10 51 62 4o

1 4n-86-46 37 45

1 48-86-78 68 63 35

1 2-5A-86-45 51 29

1 3-85-114 54 35

2 18-86-29 18 28 18 33

2 2-85-058 20 24 34 13 13

2 1-85-050 34 32 so 32 28

2 2A-86—57 42 34

2 18-86-26 27

2 3a-86—43 31 22 19 21

2 3-85-045 14 28 29 27 25 22 21 27 so
* 1 = flxzus.monandaez 2 = Cinania sp.: 3 = bacnoslebum

eunbonbiae

fifliiflll 6 = Sltbblumlaxenae: 7 =

: 4 = Eucenanbls.sv- and uzzus sp.: 5 = nxzus
Dastxnotus.

 

159

Table 44.. (cont’d.)

 

Aphids per Cell (cell 1 inner-most)

 

Aphid Trapnest

 

spp.* number 1 2 3 4 5 6 7 8 9 10
2 28-86-51 51 29
2 28-86-02 26 29 28
3 2-85-078 33 43 38 25
3 38-86-27 29 30 20 32
3 38-86-78 29 38 27 25
3 2-58-86-50 21 18
3 2—58-86-18 28 9 16 28 17 16
3 3A-86-80 22 23 28 17 20 19
3 48-86—57 26 15 17
3 2-58-86-19 13 13 13 18
3 3-85-006 13 13 11
3 2-58-85-65 9
3 3-68-86-35 24
3 3-85-075 14
4 1A-86-47 28 18 20 20
4 28-86-26 39 33
4 28-86-15 37 33
5 1-85-032 3 37 23 32 27
5 28-86-21 69
5 3-68—86-17 74 39 43
6 1-85-059 27 24 46 29 47 36
7 2-85-052 27 30
? 3—85-085 20 25 26
? 3A-86-07 41 43
? 3-85-029 36 76 63 44 48 62 58 43
? 4-85-003 75 71 86 55 47
? 28-86-15 32 37 36 38

Totals 2928 2521 1608 1065 686 411 225 93 56 25

 

*1=111Lzusmonandae:2=mnar.i.asp.z 3=ummgm

 

160

Table 45.

 

Trap Nest Data on Aphids provisioned by Eassalgggu§_guseidatu§, 1987.

 

 

 

Aphid Trap Nest Total Avg. 3 of t of days date
spp.! number in nest per cell cells worked closed
1 1-85-116 95.00 31.67 3 7 6-12-87
1 2-86-18 53.00 53.00 1 2 6-13-87
1 1A-86-50 44.00 44.00 1 2 6-13-87
1 2A-86-24 73.00 36.50 2 3 6-15-87
1 3-86-85 108.00 36.00 3 4 6-15-87
1 2-85-096 83.00 27.67 3 3 6-15-87
1 2—85-083 74.00 37.00 2 4 6-18-87
1 4—85-049 128.00 42.67 3 4 6-18-87
1 3-6A-86-15 106.00 26.50 4 4 6-18-87
1 18—86-14 64.00 21.33 3 2 6-19-87
1 5-85-115 95.00 47.50 2 7 6-19-87
1 28-86-66 111.00 27.75 4 3 6-20-87
1 2-5A-86—62 192.00 32.00 6 0 6-24-87
1 3—6A-86-46 238.00 47.60 5 5 6-24-87
1 2-58-86-44 147.00 29.40 5 6 6-25-87
1 6-7A-86-15 87.00 43.50 2 3 6-26-87
1 3-68—86-67 167.00 33.40 5 6 7-01-87
1 2-5A-86-81 192.00 32.00 6 7 7-01-87
1 2A-86-12 201.00 50.25 4 2 7-03-87
1 3-68-86-14 202.00 20.20 0 3 7-06-87
1 3A-86-58 29.00 29.00 1 2 7-05-87
1 2-5A-86P68 222.00 31.71 7 8 7-05-87
1 2-85-055 199.00 39.80 5 2 7-05-87
1 38-86-79 274.00 45.67 6 2 7-08-87

 

 

*
...
ll

llmsmnandae32=flnamsoz 3=iiaszroslobun
euobonbiaes4=mcecaobisso andlimsso45=lixzu§
maiz=suob1umamae37=nactmfl

161

Table 45.. (cont’d)

 

Aphid Trap Nest
spp.* number

 

2-5A-86-67
2A-86-44
4A-86-31
28-86-32
38-86-21
35-86-47
25-86-27
3A-86-85
3-6A-86-66
2—53-86-07
2-5A-86-72
3-68-86-20
28-86-82
3A-86-32
3A-86-60
3-68-86-07
45-86-68
3-68-86-10
48-86-46
48-86-78
2-5A-86-45
3-85-114
18-86-29
2-85-058
1-85-050
2A-86-57
18-86-26
3A-86-43
3-85-045

NNNNNNNH".”O‘HHHHHHhfithHh-Hpuppp.‘

Total
1n nest

205.00
51.00
108.00
78.00
116.00
236.00
42.00
187.00
74.00
87.00
89.00
165.00
75.00
326.00
141.00
100.00
100.00
153.00
82.00
158.00
80.00
89.00
97.00
104.00
156.00
76.00
27.00
93.00
223.00

Avg.

per cell

41.00
25.50
36.00
39.00
38.67
59.00
42.00
31.17
74.00
29.00
44.50
23.57

48383834
8388888818

3 of
cells

 

~0bfi‘NCflmANNQNQNNQQN‘lNGt-eGHAwNQNOI

4 of days
worked

mwflflAHANQmQQGmmmmafimmm-hNNHNNsl

date
closed

7-01-87
7-08-87
7-08-87
7-08-87
7-10-87
7-10-87
7-14-87
7-14-87
7-14-87
7-14-87
7-14-87
7-14-87
7-20-87
7-20-87
7-20-87
7-20-87
7-20-87
7-20-87
7-20-87
7-20-87
7-22-87
7-24-87
6-02-87
6-03-87
6e06-87
6-07-87
6-08-87
6-08-87
6-08-87

 

*1alixzusmonandae12=mnar_i.aso-:
41121192111119.44sz andMusso.:5=llxzua

m;6=mmzv=m.

3=mmnhm

 

162

 

 

 

 

Table 45., (cont’d)

Aphid Trap Nest Total Avg. 4 of 4 of days date

spp.* number In nest per cell cells worked closed
2 2A-86-51 80.00 40.00 2 1 6-13-87
2 28-86-02 83.00 27.67 3 4 6-12-87
3 2-85-078 139.00 34.75 4 2 7-05-87
3 3A-86-27 111.00 27.75 4 7 7-08-87
3 38-86-78 119.00 29.75 4 4 7-08-87
3 2-58-86-50 39.00 19.50 2 2 7-10-87
3 2-58-86—18 114.00 19.00 6 6 7-10-87
3 3A-86-80 129.00 21.50 6 4 7-14—87
3 48-86—57 58.00 19.33 3 6 7-20-87
3 2-5A-86-19 57.00 14.25 4 8 7-22-87
3 3-85-006 37.00 12.33 3 7 7-27-87
3 2-5A-85-65 9.00 9.00 1 5 7-29-87
3 3-6A-86-35 24.00 24.00 1 - -
3 3—85-075 14.00 14.00 1 10 8~06~87
4 1A—86-47 86.00 21.50 ’4 5 6-03-87
4 28-86-26 72.00 36.00 2 5 6~03-87
4 2A-86-15 70.00 35.00 2 4 6-06-87
5 1-85-032 122.00 24.40 5 21 7-05-87
5 28—86-21 69.00 69.00 1 3 7-08-87
5 3-6A-86-17 156.00 52.00 3 2 7-10-87
6 1-85-059 209.00 34.83 6 10 6~08~87
7 2-85-052 57.00 28.50 2 5 7-29-87
? 3-85-085 71.00 23.67 3 1 6~08-87
? 3A-86-07 84.00 42.00 2 5 6-19-87
? 3-85-029 430.00 53.75 8 8 7-14-87
? 4-85-003 334.00 66.80 5 4 7-14-87
? 28-86-15 143.00 35.75 4 3 7-27-87

Totals 9618.00 34.23 281 381
HWWZ= QinarJaso: 3=11aszro512hum

93133136

W14-Wso.andmso:5=m
=3119b1uma1enaes7=nactxnotus

 

APPENDW 5

Proposed Future Investigations

Prior to thls investigation 2am 3119111118 was reported
from only two 1.5 mm bore trap nests (Vlncent. 1978). My studies have
identified an area with a slgniflcant population of E. W and
suggest factors significant in its distribution. To verify and expand
upon these studies. 1t is proposed that a study of the following
aspects of the trap-nesting blology of W W daring
the sumer of 1990 be undertaken: preferences for station species,
bore diameters, and nest height: mration of the provisionlng season,
and species of aphids provlsioned.

Pre-split paraffin-coated trap nests with bore dlameters 1.2 -
2.8 mm wlth increments of .4 mm will be drilled to a depth of 60 mm.
Fifteen statlons will be selected, including five each of Minna,
Bing: and Enaxlnus. Trap nest bundles will be dlstrlbuted at heights
of one to slx meters with one meter intervals. Bundles will consist
of fifteen trap nests - three from each of five bore dlameters.

Trap nests will be distributed in the study area in mid-Nay.
Inventories will be made each week. closed nests will be removed and
replaced with trap nests the same bore diameter. Closed nests will be

opened. data taken on nest architecture and provislons, and two aphids

163

‘-

 

 

164

will be removed from each provisioned cell for Identification.
Observatlons will be continued through mid-August. Wasp larvae and
provlsions will be transferred to rearlng vials and stored over winter
In an unheated garage. Adults reared the following spring will be
used to confirm Identifications.

Chi-square (I) will be used to test for dlfferences In
preferences for station species. bore diameters and trap nest heights.
One way ANOVA and t(II) tests will be used to test for differences in
cell lengths and volumes among trap nests of different bore dlameters.
Aphid provislons will be analysed for dlfferences in numbers of aphids
provlsloned and camared with aphids provisioned by 2. cumidatus to

determine whether these wasps partitlon aphids by species.

Q

LITERATURE CITED

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Chrysldidae in America north of Mexico. Mem. Amer. Ent.
Inst., no. 33.

Bohart. R. M. and A. S. Menke. 1976. Sphecid wasps of the
World. Ix + 695 pp. University of California Press.

Corbet. S. A.. and M. Backhouse. 1975. Aphid Hunting Wasps:
A Field Study of Eassalgggus. Tran. Roy. Ent. Soc.
London. 127: 11-30.

Danks, J. V. 1971. Biology of some stemrnesting aculeate
Hymenoptera. Trans. Roy. Ent. Soc. London.
122: 323-399.

Dixon, A. F. G. 1973. Biology of Aphids. The Institute of
Biology’s Studies in Biology no. 44, 58 pp. Edward Arnold
(Publishers) leited. London. '

Fye, R. B. 1965a. Methods of placing wasp trapnests in
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._____ 1965b. The Biology of the Vespidae, Pompilidae and
Sphecidae from trapnests In northwestern Ontario.
Canadian Ent. 97: 716—744.

Cause, 6. F. 1934. The Struggle for Existance. Vllliam 8
Nllklns, Baltimore. Reprlnted 1964. Hafner Publ. Co.,
New York. 163 pg.

Gause, G. F. 1935. Behavlor of Mlxed Populations and the
problem of natural selection. Amer. Natur. 69: 596-609.

Krombein, K. V. 1955. Miscellaneous prey records of solitary
wasps. 1. Bull. Brooklyn Ent. Soc. 50: 13-17.

1956. Miscellaneous prey records of solitary wasps.
11. Bull. Brooklyn Ent. Soc. 51: 42-44.

1958. Miscellaneous prey records of solitary wasps.
III. Proc. 8101. Soc. Washington. 71: 21-26.

1960. Biological notes on some Hymenoptera that nest
in sumach pith. Ent. News. 71: 29-36. 63-69.

165

 

 

 

166

_____. 1961a. Miscellaneous prey records of solitary wasps.
IV. Bull. Brooklyn Ent. Soc. 56: 62-65.

. 1961b. Eassalnecustuninnum Dahlbom. an
adventive European wasp In the United States. Ent.

News. 72: 258-259.

 

_____ 1963. Natural history of Plummers Island Maryland.
XVII. Annotated list of the wasps. Pro. Biol. Soc.
Washington. 76: 255-280.

. 1967. Trap-nesting wasps and bees: life histories,
nests. and associates. iii-vi + 570 pp. Smithsonian
Press, Washington D.C.

 

Muesebeck, C. F. W., K. V. Krombein. et. al. 1979.
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Park, T. 1948. Experimental studies of Interspecies
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beetles. 12111111111111 992mm Duval and W
W Herbst. Ecol. Monogr. 18: 265-308.

Smith, Ray F., L. P. Anderson. and N. T. Reynolds. 1950.
Insect Pests of Alfalfa seed. California Agric.
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Smith, Ray F., W. M. Hoskins, and 0. H. Fuilmer. 1948.
Secretion of DDT in milk of dairy cows fed low-residue
hay. Jour. Econ. Ent. 41: 761-764.

Smith. Ray F., and A. E. Michelbacher. 1949. The Development
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in foot hill areas of California. Ann. Ent. Soc. America
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Vincent, D. L. and J. D. Hoffman. 1974. Advantages of using
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Yarrow. I. ll. H. 1970. Some nomenclatorial problems in the
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