Wasp Studies

3P
S T U DI ES
AFIELD
B!
H I L R AU AN D N E LLI E R AU
W I T H A N I N T R O DU C T I O N
B!
W I LLI AM M W H E E LE R
.
N O M IC E N T O M O L O G Y
HA RVAR D U N I VE R S I T Y
P R O F ES S O R O F E
P R I NCE T
VE RS I T ! P R E S S
P R I N CE T O N
LON DO N : HU M P H R E ! M I LFOR D
O XFOR D U N I VE R S I T! P RE S S
ON
UN I
1 918
P I
R
Pi
C o p y ri g ht , 19 18 , b y
N C E T O N U N VE R S I T !
r n te d
ub
l i sh e d
c t ob e r ,
i n t he U n i t e d
19 18
t a te s
of
R ES S
Am
eri c a
Psy cholo gy is an as y e t almost u n explored

terri tory a n d but li ttle u n de rsto o d for wa nt of appro ach in g
i t b y th e best si de th a t is to s ay b y c a refully m a de o b s er
v a ti o ns
I t is inv olved e i th er i n m etaph ys i c a l do g mas
or i n sha llo w a n throp omorph ism w h ich co n fo un ds i h
heri ted i nst in ct a n d i ts a uto m a t isms wi th th e pl asti c ju dg
me n t of the in divi du al b ased up o n me m ory a n d the as so
c i a ti on of m emori es or se n sory i mpress i o n s
L et us
b e th oro ug hly imb ue d wi th the tru th tha t e a ch sp ec i es a n d
ev en e a ch p oly morphi c a n im a l form ha s i ts spec ial p s y
c hol og y wh i c h should b e especi a ll
stud
i
ed
w
h
i
ch
de
n
d
a
y
p e n ds o u the on e ha n d up o n the dev elopme n t of i ts muscles
a n d se n ses a nd on the o ther up o n tha t

n
i
ts
br
a
i
of
F O R E L The Sense s o f I n s ec ts
Co m p ar a tive
P REFACE
The present volu m e em

bodies the results o f four
rs o f out o f door study
e
a
y
o f some o f our most in
teres t in g and hi ghly de
insects in thei r
v el o p ed
native haunts while pur
suing thei r occupations in
thei r own w ay B io logical
behavior work on the Amer
ican wasp s has been fo r the most
part desulto ry and incomplete and
hope that these chapters m ay in thei r small
w a y ll the ga p that exi sts
We have no ap o logy to make for the f re
quent use o f anthropomorphi c ideas terms and
interpretations H owever one must n ot read
into these terms any subtle meta p hysical mean
ing
They are used a s apt descriptive ex
pressions and not for the purpose o f predi
cating logical thinking to these creatures
Thi s work as a whole is descriptive ; the
comparative and philosop h i cal a s well a s the
co rrelative data have been reserv ed for a later volume
Unl ess otherwise stated all o f the o bservation swe re made
at S t Loui s or w ith in a radiu s o f thi rty miles o f that city
The sketches for the illustrations were made in the eld and
executed in the i r nal form by D r G ustave Dahms
-
v ii
A debt o f gratitude i s due to D r C H Turner o f S um

ner H igh S chool o f thi s city fo r critical ly reading the
manuscript and to D r L O Howard and the various spe
c i a l i s t s connected w ith the United States Department o f
Agricultur e and the Smithsonian Institution whose names
appea r i n brackets throughout thi s work w e desi re to ex
press our thanks for the i denticati o n o f the va rious insects
and especially i s our gratitude due to M r S A Rohwe r fo r
much taxonom ic in fo rmation
P H I L R AU
N E LL I E
S A I NT L O U I S Mo
November 2 9 1 9 1 7
.
viii
CO NTE NT S
I N T RO D U CT I O N
W M W H E E LE R
By
C H AP T ER I
Some
B em b i c i n e
Wasps
B emb ix n ubi lip en n is

B e m b ix s p i n ola e
M ic ro b em b i x
mo n odo nta
B i cy r tes q ua drif as c i a ta
Sphec ius s p ec i os us
'
C H APTER I I
B ehavior
o f Wasps
B elonging t o the Fam ily P omp ilida e
45
P o mp i l oi d es tr op i c us
P omp i l oi d es
ma rgina tus
P s a mm o c hares
s c el es tus
P epsis dub i ta ta
p omp ilus
Arac hn op hoc to n us ferrugin eus
P ri o c n emis
P s eud ag en i a
a rc hi tec ta
P s eud ag en i a p u l c hrip en n is
P s eud ag en i a mellip es
C H APTER
S ome Fly catch i ng Wasps
-
0
9
H yp oc r a b r o
s ti r
H yp ocra b r o
c hrys arg i n us
i
o
l
u
c
s
p
P ara n o thy r eus
III
c i n g ul a tis
ix
C HAPTER IV
The Enemi es o f P lant lice
-
1 02
Xy l oc eli a m e ta thorac i c us
C era top horus t en ax

C HAP T ER V
The
B ee- kill i ng
Wasps
1 09
P h il a n thus p un c ta tus
P s euda n thop hi lus
ver ti la b ris
C HAPTER VI
Some Mud daubing Wa sps that H unt Spiders
S cel iph ro n caem en tari um
C haly bi on c a eru l eu m
-
1 1
C HAPTER VI I
S ome Wa s ps that
Cerc eris
P rey
on
B eetles
1 26
um ip enn i s
S colia dub ia
C H APTER VI I I
S ome Wood boring Wasps
C erc eris n i ti ma
Tryp oxy l on tr id enta tum
S i la on n ig er
-
Tryp oxy l on
cla va tum
Tryp oxy l on
33
0
4
o hns on i
C HAPTER I X
The H unters o f Small O rthoptera
A ly son m ell eus
Tac hys p hex termi n a tus
Tac hy s p hex
Tac hys p hex
N o tog on id ea
us c us
o b s c urus ,
and
o rg en ta ta
o b duc tus
'
C HAP T ER X
T he
H unters
Large O rthoptera
of
P ri on o n y x
P ri on ony x
T he
59
tra tum
thoma e
of
relation
S tie us
un i c i nc tus to
P ri on on yx
th omae
Chl ori on i c hne um one um
A locust hunt er that makes twin cells

Chl ori on cy an eum
Chlo ri o n ( I s o d on ti o ) Aurip es
T he prey o f Am m o bi a b ri dw elli
-
'
'
C H AP T ER X I
T he
Sand loving Ammophila

Sph ex p i c tip en nis
Sph ex proc er a
2 07
C HAPTE R XI I
Experiments o n
Some S oc i al Wasps
P ol istes p a llip es
P olistes ru b igi nosus
P ol is tes an n ular is
P ol istes p allipes
P olistes b el li c os us
Ves p ul a g erma nic a
Vesp a mac ula t a
.
the
H oming
of
2 44
'
T he
C HAP T ER XI II
M in ing a n d Other Wa sps o f the Family Eumenidae
O dy nerus g em in us
O dy nerus dorsal is
O dy nerus an orm is
O dy n er us p ed es tris
O dy nerus d esig na tis
O dy n erus f ora m i na tus
xi
2 97
An ci s troc erus f ulvi p es

An cis troc er us t ig r is
A n c is tr oc erus
co
ra
M o n o b i a q ua dri d ens
C H AP T E R X I V
G ene ral Considerations
Index
xii
35 5
ILLU STRATI O N S
F IG
I
Nest o f the p otte r wasp E u me n es s p

An attractive area for digger wasps
The burrow s o f B em b ix n ub i lip en n is
B emb ix n ub i lip en n is ll ing her burrow
Diagram S h o wi ng the meth o d o f B em b ix
pe n n is i n sweeping the ground
-
7
8
10
1 1
12
28
n
ub il i
1
3
The cocoons o f B emb ix n ubi lip en n is

The burrow o f M i c ro b em bix mo n odo n t a
A sandy area where M i c r o b em b ix mo n odo n t a and
B i c y r tes q ua dri f as c i a ta a b ound
P o m p i l o i d es trop i c us transporting her prey
The burrow o f P o m p i l oi d es trop i c us
Another f o rm o f P o m p i lo i d es trop i c us burr o w
P o mp i l oi d es m a rgi na t us dragging her prey up hill

The burrow and p rey o f P s a m m oc ha res s c el es tus
P ri o c n em i s pomp ilus carrying her spider
Figure o f female Arac hn op ho c to n us ferrugin eus
The twin celled nests o f P s eud ag en i a arc hi tec ta
P s eudag en ia p ul c hr ip en n is and her prey
The nests o f P s eudag en i a m ellip es i n the walls o f
a S celiphron nest
A three celled nest o f P s eud ag en i a m ellip es and
its make r
The mud cell s o f P s eudag en i a from which
S p ha er op h thal m a s c a ev a emerged
A mud nest o f P m ellip es built i n an oak apple
The burrow a nd cells o f H yp ocra b r o s ti rp i c o lus
.
35
4o
43
49
53
6
5
61
65
6
7
79
84
85
86
! OA
! OB
21
21
IO
7
18
PA G E
87
xi i
88
89
22
The remains o f a feast ; cell s o f H yp oc ra b ro sti r

p icola s
An elde rberry twig containing the nest o f H yp o
23
cra bro
24
25
2
9
i
u
c
o
l
s
p
s ti r
95
The nest i ng sit e o f P aran o thy r eus c i ng ula ti s

B urrows o f va r i ous fo rms made b y P a ran o thyr eus
97
c i ng ul a tis
26
27
28
29
99
Figure o f female P a ra n o thy reus c i ng ula tis

A clay bank wh ere Xy l oc eli a m eta thora ci c us digs
her bu rrows
The burrow and cells o f X y l o c elia m eta thorac i c us
The burr o w o f P h il an thus p un c ta tus
N ests 0 f the mud da ub er 5 c elip hron c a em en ta r
i um and C ha ly b i on c a eruleum
Th e opening to the burrow o f C erc er is f um ip enn is
A S c el iphro n nest remodelled to su i t the needs o f
1 01
1 03
1 06
1 1
1
3
2
3
Tryp ox y l o n
clava tum
1 20
1 26
6
3
33
A ly so n m ell eus ,
he r burro w and p rey
1
4
34
The bu rrow o f
Tac hys p hex ter m i na tus
6
4
6
5
60
63
64
35
37
8
3
39
0
4
1
4
2
4
43
44
45
46
47
8
4
rg ent a ta and he r burrow

P ri on ony x a tra tum
P ri on onyx a tr a tum ll ing her burrow
P ri on ony x a tr a tum po unding th e so i l w i th he r head
T w o burrow s o i P ri on o n yx a tra tum
P ri on on y x thoma e riding he r l o cust home
P ri on on y x thom a e and S t iens um c m c tus
Cinders carried by P ri on onyx thom a e
N o tog o n i d ea
C hl o ri on i c hn eum on eum he r bur row and pr ey

Tw i n celled burrows made b y an unknown loc h st
huntress
The burrows o f Sphex p i c tip en n is
Sphex p i c tip en n is carrying a small caterpi llar
The position o f egg o f Sphex p i c tip en n is on p rey
A local ity study by Sphex p i c tip en nis
,
77
1 83
85
9S
xiv
99
2 08
I
210
,
21
2 20
49
0
5
2
5
53
The cocoon s o f Sphex p i c tip en n is

A mmoph il a proc er a aslee p on a dried stem
The nest o f P olistes
A nest o f P olistes a nnula ris among the branche s
Vespula g erma n ic a thick as ies on a freshly re
moved rooster s head
The turret over the bu rrow o f O dy n er us g emin us
A burrow o f O dy n erus g em in us
A coll ection o f mud p ellets carried out o f the bur
row by O dy n erus g em in us
Five burrows made by O dy n er us g em in us
O dyn erus dorsa l is her burrow and he r method o f
loweri ng the p rey
The cours e o f ight o f O dy n erus dorsa lis to her
water puddle
The burrow o f O dy n erus dorsal is
Two burrows o f O dy n erus dorsal is
An unusual O dy nerus n est
The mandible marks on the tunnel le ft by Ody
n erus dors a l is
All that rema i ned a fter 0 dorsa lis f ea st on P h oli
sor a c a tullus caterpillars
The nest o f O dy n erus p ed es tris
O dy n erus f ora m i na tus and he r home i n a log
The nest o f O dy nerus f ora m i na tus in an elderberry
stem
The burrows o f a min i ng bee in a clay bank some
o f the tunnel s used by An c is tr oc erus f ulvip es
.
54
55
6
5
57
8
5
59
61
62
63
65
66
67
68
243
28 7
2 89
2 96
00
3
0
1
3
60
64
2 35
0
3 3
0
8
3
1
3
1
3
1
3
1
3
2
1
3
2
2
3
2
3 7
333
335
8
33
XV
2
34
W AS P
A F I E LD
S T UD I E S
I N TRODU CTI O N
I f any excuse were needed for welcoming another work
i n additi on to the nearly three hundred books and articles
that have been publ i shed on the habits o f the sol itary wasps
it would su fce to po int to the fact that no other grou p o f
i nsects has s o fasci nated and b a i ed the student o f animal
behavio r the psychol o gi st and the philosopher
When
among contemporary general izers we nd an eminent
p sychologist William M c Do uga ll claiming for the sol itary
wasp s a degree o f i ntelligence which ( w ith the doubt ful

exception o f the higher mammals ) appr o aches most nearly
to the human and the illustrious B e rgson using the same
i nsects as paradigms o f instinct i n the sense o f intuiti on
as contrasted w ith intell igence there i s su rely need o f a

renewed presentation o f facts al ready e stabl ished o f the
publication o f new observations and o f a seri o us attempt
at di spassionate interpretation like that made i n the present
volume
The sol itary wasps c o mpri se some
described species
scattered over the torrid and tempe rate regions o f the globe
and rep resenting a number o f more or less closely related
natural famil ies o f Hymenoptera To the entom o logi st these
wasps are o f unusual interest fo r several reasons Fi rst they
are i n physical structure the most superbly special ized o f i n
sects so that they bear to creature s l ike the beetles ies and
grasshoppers somewhat the same relation that the members
o f the cat family bear to the rodents ruminants and i n s ec t i
vo res
Even the social Hymenoptera seem to have a less
,
WA S P S TUDI E S AFI ELD
per fected nervous and muscular organization a ri d are cer

ta i n ly less beauti fully formed and colored
S econd th is
unusual physical endowment i s correlated as woul d be ex
t
e
c
e
d
w
ith
extraordinary
i
ndustries
o
r
behavio
r
Cer
p
ta i n ly w ith the s ingle exception o f the social H ymenoptera
and the Termites n o insects S how such a range o f a c tiv i
ti es a s the sol itary wasp s Third they are the l ineal de
s c en da n t s o f forms wh ich gave ri se to the social H m en o p
y
tera Thi s seems to b e s o evi dent that H a n dl irs c h actually
derives the bees fro m the Sphegid the soc ial wasps f rom
the Eumen i d and th e ants f rom the S col i i d wasps and
R o ub a ud has recently been able to detect i n th e genera Sy n a
g ris an d B elon og as ter o f the B elgian Cong o a most inte r
esting series o f behaviori s tic trans itions between the sol i
tary and social wasps And fourth the ancestry o f the
sol itary species themselves presents an interesting though
more d ebatabl e p roblem owing to the fact that the group
appears fully developed at least s o far a s the famil ies M util
l id ae and S col iid ae a re concerned i n the B alti c Amber o f
lower Tertiary age Although no species have been found
i n olde r geol o gical formations we must sup p ose n ev erthe
less that the group go e s back to the Cretaceous and prob
ably even to th e Jurassi c or Triass ic Hence the evoluti on
o f the sol itary wasp s has extended over a period o f at least
four to s ix mill ion years We are n ot there fore greatly
surprised to nd that they exhibit such a d ivers ity o f hab its
especially when w e remember that the whole mammal ian
class man o f course included has had a shorter evolution
There i s anothe r pecul iarity o f the solitary wasps which
i s connected with thei r geologic al o rigin and h istory The
autho rs o f th is book repeatedly call attention especially i n
thei r account o f the B em b i c id s ( Chapter I ) to the fact
that these i nsects a re very strongly attached to thei r nesting
s ites and stick to them generation a fter gene ration Such
.
o f the complete cycl e o f behavior o f any single species can

not be thus obtained Field study is there fore indispensabl e
and thi s demands acute powe rs o f observation much p a
t i en c e and no l ittl e me re phys ical endurance
S ti ll the
l im i tations o f such study are obv i ous The nding o f the
insects and the i r nests i s a matter o f chance and frequently
only gl impses o f thei r behavior are obtainable s o that the
whol e cycle o f behavio r has to be p i eced togethe r from
f ragmentary observations a n d o ften requi res the labo r o f
several observers extending ove r many seasons H ence the
i ncompleteness o f many accounts i n the l iterature A good
exampl e o f thi s l im itation i s see n in the p resent volume in
the interesting account o f the extraordinary parasitic habits
o f S tie n s un i c i nc tus
Anothe r l imitati o n i n eld study l ies i n one s inab il ity
to estimate the p revious experience o f the wasps unde r o b
se rvation Usually the ins ects are rst encountered while
engaged in some activity which may or may not have been
repeated on some former occasion such as making the nest
or p rovi sioning it and a s they undoubtedly pro t by ex
i
and
form
d
n i te habits
what
i
s
evidently
an
n
r
e
e
c
e
e
p
important factor in the pe r fo rmance o f the particular a c tiv
i ty unde r observation cannot be taken into consideration
Thi s lacuna i s partially b ri dged in Chapte r XI I o f the pres
ent book by a valuabl e study o f individual exper i enc e in one
o f the primitive wasps P olistes
Even more seri ous i s the l imitation to the experimental
method in the el d I t i s o f course essential fo r d es c rip
tive and other purposes to ascertain by s imple observation
the course o f the normal routine activities o f the insect i n
i ts natural envi ronment but i t is also o f great im p o rtance
to know what the insect i s and i s not capable o f do ing Thi s
can be accompl i shed only by means o f e xperiment ; and
experimentation i n the el d i s a rathe r mild procedure
.
I NTRODU CTIO N
compared with what can be accompl ished unde r the rigorous

and indenitely vari able control o f the laboratory Still
results o f considerable value have been obtained by simple
eld experimentation as will be apparent to the reade r o f
the ingenious studies o f Fabre the P eckhams and the Raus
Although the activities o f only a few hundred sol itary
wasps have been care fully observed w e may be sure that
eve ry one o f the
described species has its own p ec u
l iar behavior I n the non parasitic f o rms thi s appears as
a complex cycle the more impo rtant component m inor
cycles o r phase s o f which are the digging o r construction
o f the nest the capture and stinging o f the insect or sp ider
prey ovipos iti on and the seal ing o f the nest entrance B ut
the sequence and details o f thes e cy c l es i s subj ect to great
sp eci c and sometimes to considerable individual variation
Thus the sequence o f the three rst cycl es i n many speci es
o f S p heg i d s is nest
p rey egg but i n P s a m m o c ha rid s it
i s commonly prey nest egg and in the E um en id s nest

eggp rey Thi s i s al so the sequence in social wasp s ( P o
l istes Vesp a
I n parasiti c species the behavio r is
o f c o urse peculiarly modi ed i n adaptatio n to that o f the
host
I n the i r interp retati on o f wasp behavio r the Raus agree
essentially w ith nearly all p rev ious i nvestigators as could
be shown by quotation s f rom Marchal P icard B ordage
Adle rz the P eckhams H artman and others Most o f the
activitie s can b e readily interpreted as chain re ex es or
insti ncts i n the usual b i ological se ns e o f the term They

are relatively xed or stereotyped and undoubtedly heredi
tary and there fore rep resent the most ancient and mos t
B ut there
sol idied complex o f the behavio ri sti c cycle
stand out from thi s complex many activitie s wh ich are much
l ess mechanized and o f such a nature as to demonstrate
that the wasps p o s s es s s emotions and associative memory
.
'

-
that they exercise discriminati on and choice that they learn

by experi ence and form hab its i n the restricted sense o f the
term and that they can modi fy thei r behavio r adaptively in
response to unusual stimul i on the basi s o f previous ex p er
i en c e and there fore behave to a l imited extent l ike i n telli
gent beings
Such a n i nterpretation will be accepted by
any unbiassed student o f the sol itary wasp s and those
who hold i t may be classed as the moderates among
behavi ori sts
There are however three classes o f extremi sts the
mystery mongers the s i m p li c i s ts and the humanizersw li o
fai l to take accou nt o f a ll the facts and nd support f o r
the ir o p inions i n particular aspects o f wasp behav ior
Fabre may be classed among the mystery monge rs because
when he turned to the i nterpretation o f the wonder ful facts
he had s o ca re fully obse rved hi s nal a ppeal w a s always to
some mysterious cause Thi s i s bes t seen i n hi s interpreta
tion o f the stinging o f the p rey as the exp ression o f an in
scrutable i ntuitive knowledge o f the prey s nervous a n
atomy a v i ew wh ich B ergson i nco rporated i n hi s p hilo s o

phy o f instinct Fabre s treatment o f the wasp s hom ing
activities and o f her method o f nding he r p rey are sim ilarly
attributed to he r pos sess ion o f mysterious senses
Thi s
mental attitude w a s due to p reconceptions to w hich I shall
return pr esently The s im p l ic i s ts are p reoccupi ed w i th the
reex o r trop i stic i e the mechan ized behav ior At least
one o f them B ethe i s also as much o f a mystery monger
a s Fabre Loeb and B o h n have no rst hand acquaintance
w ith the wasp s and are inuenced by thei r knowledge o f
the l o we r i nverteb rates and insects whose behavior i s s o
highly and cons istently mechan ized that evidence o f its
plasticity or m o d i ab ility are feeble or di fcul t to observe
Th e humanizers who i nterpret animal s l ike the wasps as
i f they we re mi niature men are rarely met w ith at the
,
'
I NTRODU CTIO N
present time except among the writers o f certai n nature

books
The case o f Fabre requires some further comment Ac
h
s
n
n
i
a
c
e
o
are
not
entomol
gists
but
who
read
hi
s
u
a
t
w
o
q
Souveni rs w ith del ight frequently ask whether I regard

him as a rel iabl e chronicle r o f the habits o f insects
I
usually rep ly that in my opin ion Fabre Latreille and R eau
mur are the three greatest entomologists but that we must
make due allowance i n Fabre s writings fo r certain p rec o n
c ep t i o n s which strangely en o ugh seem actually to height
the merit and beauty o f h is work H e wa s trai ned as a
physici st chemi st and mathematician and when he came to
study i nsects he carried w ith him the po int o f view o f the
student o f the ino rgani c science s and was there f o re inter
e s ted in establ ishing clean cut laws
Ow ing to this tendency
and a bel ie f i n the scholasti c conception o f instinct he natu
rally stressed and schematiz ed the normal course of behav
i or i n the insects Although he noted many va riations in
thei r activity these evi dently appeared to him as s o many
erturbations
o
r
accidents
which
w
ere
interesting
only
in
so
p
far as they helped to de ne hi s essentially static interp reta
tion They were i n othe r words me rely the exceptions
that prove the rul e Hence hi s descriptions and discussions
leave an impression o f elegance and nal ity like a demon
s tra ti o n i n mathematics o r phys ics and there fore ap p eal
more strongly to philosophers like B e rgson than to t he mod
ern biologist w ho has been so o ften deceived by clean cut
theories co nce rn ing l iving organi sm s that he has grown
tim id and susp icious
Then i t must be rememb ered tha t
Fabre w a s unable to app reciate the variati o ns o f behavior
because he was to o s et in hi s ways o f thinki ng when the
Origin o f Species appeared to acquire any sympathy w ith

evolutionary theories I f these considerations be borne in
m ind i t i s not di fcult t o estimate the value o f Fabre s
,
work
H e i s indeed so p reeminent in the wealth and pre

in the ingenuity o f hi s ex p eri
c i s i o n o f hi s obser v ations
mentation and in l iterary expression that h i s Souven irs

w ill always endure
The variation s which to Fabre were more o r less negl i
i
l
necessarily
at
once
assumed
great
importance
when
b
e
g
biol o gi sts became evolutionists I n fact the variations in
behavior because they were consi dered to b e the essential
material s o f evolution became i f anything even more in
Th e rst
t ere s t i n g than the routine m echanized activities
among entomologists to recognize the evolutionary impor
tanc e o f behaviori sti c variati o ns was P aul Marchal i n 1 88 7
i n h i s study o f a sol itary wasp C er c eris or n a t a and s ince
that time all students o f insect behavior have focussed thei r
attention on the variations Thi s i s a s apparent in the pres
ent volume a s i n the works o f Marchal the P eckhams
Adlerz F e rto n I seley and others and coul d not well be
otherw ise fo r all biologi sts are now thoroughgoing evo lu
.
t io n i s ts
In one othe r respect al so the Raus have followed a com

mendable tradition i n the presentati on o f thei r observations
Re aumur in hi s M emoi rs publ ished i n 1 734 to 1 74 2 w a s

one o f the rst entomol ogists t o w rite on the behav i or o f
insects in such a manner as to attract and i nstruct the gen
eral reade r Fabre adopted a S im ilar method o f p resentation
but greatly excelled hi s ei ghteenth century p redecessor i n
l iterary p ower That the Ameri can i s as app reciative a s
the European publ i c o f this form o f writing i s shown by
the success o f the P eckhams w ork on the sol itary wasp s
I feel sure that Wasp Stud ies A eld w ill meet w ith a
l ike rec ep tio n x
W M W H EE L E R
Harvard University
June 1 0 1 9 1 8
,
CHA P TER I
SOME
B emb ix
n ub i l i
en n is
W ASP S
B E M B I CI N E
Cress [ S A Rohwer ]
.
The B emb ix population burst upon us w ith a suddenness

which startled us into full attention at once We had crossed
the eld day a fter day and on that ve ry day June 1 6 1 9 1 4
had passed by this certain bald ba re space in the eld which
the boys o f the neighborhood had for a number o f years
kept packed hard for thei r Sunday baseball game ( g 2 )
but all had then only an hour be fore been as quiet and l i fe
less as the grey earth itsel f Now the very ai r above the
sur face o f the bare ground seemed vibrant w i th the low
ying wasps which formed a wave ring yellow ish green
haze ove r the smooth dusty earth
Any estimate o f thei r numbers was ve ry di fcult to obtain
f rom the swi ftly moving swarm but we suspect that at least
one or two hundred were present The ground was dotted
1
w ith newly opened holes less than o n e fourth inch in diame
ter wh ich seemed to go straight down and had no trace o f
excavated di rt around the i r mouths ; this evidence led us
to conclude that these wasps had all s imultaneously emerged
by these exits f rom their w inter quarters to m ingle in th is
rs t social frol ic o r dance
They remained in thei r unceas
ing ight at a uni form and constant height al l keep ing w ith
.
We
c o un e d
c o ul d n o t
be
f o rty f o ur
c e r ta i n
if
a ll
ho l e s i n
e re
o n e a r ea
ho l e s
9
of
t he
t h ree
f eet
e m erg i n g
b ut w e
s q ua re ,
em b ix
WA S P STUDI E S AFI ELD
12
others although sometime s they would conclude w ith one

o f the w ild dashes and embraces j ust described
The l ivel iest part o f the dance when the participants were
most numerous occurred from 9 :30 to 1 0 a 111 B y
they were fewer in number and droning wearily along and
by 1 I o clock w heri we were obl iged to leave the eld both
the activity and numbers we re greatly reduced although we
could n ot di scern wh ither they were di sappearing
Two days late r we returned to the eld at 7 a m eage r
to wi tness the continuance o f th i s per fo rmance and learn
mo re o f its mysteries The holes w ere all open as be fore ;
but some m inutes elapsed be fore the rst d rowsy B emb ix
appeared and took up the low gl iding ight o f the day b e
fore P resently others j oined i t and by
ve were in
the ight grace fully w eav ing to and fro and occasionally
rest i ng fo r a moment on the grey earth We did not detect
whence they cam e They l imited thei r ight strictly to a
bare area which was p erhap s fteen feet i n diamete r and
did not venture out above the grass wh ich surrounded it
on al l si des
B y 8 o clock and during the hour follow ing p erhaps hal f
as many as on the rst day were on the eld pursuing the
same cha racteri stic low ight but to day there was no per
c ep t ib l e hum o f w ings n o em b rac i n g s no

waltzes and very
few o f the w ild i rtatious dashes Instead f rom time to
time many we re settl i ng on the earth and beg i nning to dig
v igorou sly In thi s digging they paid no heed to the old
hole s f rom wh ich w e supposed they had j ust emerged ; b ut
they began energetically digging new burrows each with a
broader spreading entrance and go ing down at an angle
I t seems
o f approximately 30 to 4 5 w ith the sur face
that the old hol es were merely di rect channel s o f escape
f rom the place where they had lain buried in the earth
th rough the wi nter and they did not tu rn to them at all as
,
TH E
B E M B I CE N E
WA S P S
further domiciles any more than a chick returns to its

egg shell
In the following year we took up a diligent watch early
in June fo r this p retty phenomenon
Our expectations
waned with l o ng waiting unti l we gave up all hope o f seeing
the frol i c again B ut on J uly 4 we were surprised by the
outburst o f wasps as sudden as be fore The cool months
o f May and June had p robably retarded thei r emergence
unti l th is late date Even thi s hy p othesi s seems almost in
credibl e when we cons ider that they came from the eggs
laid by many di ffe rent mothers o f the previ ous year from
June to S eptember and that even the ir nests which remai n
open unti l the larva i s ready to pupate were closed at w idely
di fferent periods ; yet we can now imagine no explanation
for the phenomenon other than that the wasp s which had
come to the i r maturity s imultaneously o r had been lying
dormant ready to emerge at the right condition s had all
responded to the dazzl ing sunsh ine and the ri sing tempera
ture o f th i s b r i ght mo rn ing and had al l dug the ir w ay
straight out into the l igh t to m ingle i n this rst social f rol ic
or dance None o f the characteristic nesting holes o f B em
b ix could be seen but the clear cut vertical holes by wh ich
they had emerged w ere even mo re numerous than i n the
p rev i ous year I t w a s very noticeable too that the group
w a s l imited to p recisely the areas where the mothers had
nested during the previous few years and where also the
Thus for gene ra
s un dance o f the last yea r had occurred
tion a fter gene ration they l i ve and reproduce i n the spot
where they are born
Strange indeed i s the constancy o f instinct in thi s danc
ing per fo rmance which they are neve r taught and for
which they have no further use ; nevertheless it occurs year
a fter year with each generation and i s 1 n all details the
same In thi s the sec ond year w e were alert to ferret out
-
the answers to the many myster 1 es which the rst year had
le ft us
On the rst day w e arrived at the concl us ion that there
In all the morning s
w a s a great p redominance o f males
dance w e s aw only three mating ights I n one o f these
cases the unite d pai r were in igh t when they were knocked
to the ground by two othe r mal es A struggle ensued i n
wh i ch all the males we re trying to gain access to the femal e
They w ere s o intent upon thi s a ffa i r that we could eas ily
get clos e enough to s ee all th is Even as w e watched more
and more mal es l e ft the ight and j oined the struggl ing
mass and strove togethe r on the ground be fore us for sev
e ral minutes or maybe i t w a s seconds So large an assem

blage we re they and s o intent up on the i r bus iness that we
could have scooped up the mass i n a hand ful and the hand
woul d have been lled to overowi ng Eventually the f e
mal e and one o f the mal es b roke away from the mass and
sped o ff on the w ing The trai n o f adm i re rs which fol
lowed them formed one o f the prettiest s ights w e have ever
w itnessed in the i nsect worl d ; the wasps extr i cated them
selves one by one fro m the heap and f ollowed the pai r in
unbroken success ion l ike the w o ol spun f rom the di sta ff in
a long sm ooth l i n e o f gl inting green and yellow pursui ng
them so rap idly that be fore the fugitive femal e had gone
fty feet they had overtaken he r again and fo rmed the
teem i ng mass around her Th i s struggle continued agai n
for several seconds until one cou l d not tel l what had become
o f the femal e ; but p resently one by one they abandoned the
con ict and returned to the dance
As w e have described for the previous year thei r ight
was low near to the ground The wasps were i n motion
fo r hours at a time never resti ng ; i f one chanced to drop
out or fall to the earth for an instant another perhaps
m istak i ng i t fo r a f emale o r fo r some other unknown
.
T H E B E M B I CE N E
WA S P S
reason would immediately po unce upon it w ith a heavy

thump and thus arouse it to return at once t o the dance
The other two pa irs that succeeded i n mating made the ir
speedy escape to avoi d rivalry This accounts f o r the w ild
dashes outs ide the area o f the dance wh ich we obse rved
in the rst year ; the escape from the crowd has become s o
necessary a part o f the act o f mating that a dash for sa fety
from the throng ha s become an integral part o f every s ug
gestion o f mating o r false mating
In the m idst o f thi s pretty play we were driven from the
eld by the human prop ri etors o f the baseball diamond and
when we s a w the clouds o f dust rise f rom the trampl ing
o f many spiked shoes it seemed inevitable that the l ittle
B emb ix would be driven from the homes o f the i r ancestors
In the a fternoo n we returned but found another s et o f ball
players and many spectators doing the i r best to exte rm inate
our pets The next day at
howeve r true to thei r
fatherland they we re the re and even more abundant than
on the rs t day although how they coul d have escaped
extermination during the terrible trampl ing w ill always be
Wa sps do not migrate far from the place i n
a marvel
which they are b o rn Thi s in a pe ace ful local ity might
S h ow only indi fference but when even a noisy American
baseball crowd cannot drive them from thei r home it proves
persistence o f a most surp rising degree Where these B em
We certainly do
bix had spent the n ight w e know not
not s ee how i t would have been possible f o r any o f them to
locate the i r forme r holes from which they had emerged o n
the trampled and dusty earth even i f they had s o desi red
They may have slept on the grass and weeds surroundin g
thi s area but in extensive studies o f the S leep o f insects i n
thi s el d we have neve r found a B emb ix asleep o n the
vegetation The bare ground p resented no newly dug holes
which they might have made f or their sleep ing quarters
,
but there were more o f the vertical emergence holes f rom

which newcomers had p robably i ssued
It w a s soon apparent that there were more females pres
ent than on the p revious mo rn ing There w a s a noticeable
reduction in the r ival ry when a mating took place A dozen
o r mo re matings occurre d in the low ight near to the
ground ; since there was l ess rivalry there was less need o f
a quick e scape and mating coul d go on w ith impun ity In
only two cases w a s there marked rivalry where the mated
pai rs were knocked to the ground but instead o f the teem
ing mass o f competito rs which we had seen the day be fore
there were only thre e or fou r pursuers
The duration o f c op ula ti o n i s brie f The time i s hard
to e stimate but it usually continues through about fty
feet o f ight The females are polyandrous and all indica
tions are that the males are polygamous One certain f e
male wh ich w e coul d easily follow on account o f her ex
We are
c ep ti o n a l colo r mated s i x times i n a b rie f pe r i od
sure f rom close observation that all o f these we re cases
o f actual matings In each case al so the mating was spon
tan eo us that i s the female w a s n o t coerced by the eage rness
o f the mal es to mate nor d id the rivalry o f the males cause
the separation Each time a fter the un i on had been e f
f ec ted the pai r gl ided near the ground for the us ual d i s
tance o f about fty feet and separated ; the female returned
at once to the dance and wh i rled i n and out until s he casually
p icked up anothe r partner when the pe r fo rmance was re
m
exactly
Thus
w
ith
o
re
females
i
n
the
eld
on
e
a
t
e
d
p
the second day the rivalry w a s much reduced and w e saw
none o f the extreme behavi or o f the males
Thi s also p o m ts to the habit o f p r1 0 r1 ty o f emergence o f
the m ales as we know occurs in othe r insects cf Sat
-
urn ii d s
2
R a u,
T ra n s Ac a d S c i
.
S t Lo ui s
.
23
: 1
78
914
TH E
B E M B I CE N E
WA S P S
At
in the a fternoon o f the second day all the excite
ment w as gone and all w a s quiet excepting a hal f dozen
or s o females which were beginn ing to dig the ir nesting
burrows Al l o f ou r a fterno o n observations S how that the
mating ights occu r only during the morning
Every morning f or ten days w e found B emb ix in the
eld in the characteristi c ight but as the days passed the
number o f those in the s un dance became less and less while
the number o f females busily burrow ing increased p ropor
t i o n a tely until July 1 4 when the dancing behavior ceased
During the latter part o f thi s period o n ly an occasional
mating was to be seen What becomes o f the males ? Do
they fertil ize the females once for all and p romptly die
o r do they hide away among the grass to be at hand when
?
needed
We suspect that the former condition holds for
we have never found B emb ix males i n the grass o r any
where except fo r thi s one occasion and I have never seen
these wasps mate while nest building
Thus it seems that the one gala day wh ich we so for
tun a tely witnessed was not only the rst but also the chie f
festivity o f the B emb ix year f or although we s a w a cer
tain amount o f thi s behavior on subsequent mornings w e
never again s a w the whole population give itsel f up to the
dance as on thi s rst occasion A fter this had waned only
a few wasps were to be found above ground at a time We
do n o t k n o w whethe r the males had migrated or perished
but w e have every reason to bel ieve that a large number o f
females were busy working in thei r underground nests or
foraging fo r food for the i r young so that now only a small
proportion o f the population at a time was i n evidence above
ground
The work o f most o f our sol itary wasps in nest building
is characterized by an admi rable constancy and zeal but
some labor w ith calmness and patience a n d some even so
-
'
'
cunn i ngly that w e can seldom spy them at thei r work B ut

there i s n o chance for doubt when B emb ix i s abroad ; others
m ay equal her i n quiet dil igence but s he works w ith un
equalled commotion and bluster S ince these are sol itary
wasps working in commun ities and have a hab it o f robb ing
each other o f p rey as we shall s ee later they must work
nervously A wholly sol itary wasp or one which works in
cooperation instead o f competition w ith othe rs can a fford
to wo rk much more calmly and del iberately S he al ights
upon a spot on the bare sur face o f the eld a s p o t w i th

out a trace o f anything whi ch w e can discern to d is tinguish
i t from all th e rest o f the large bal d area several yards in
w i dth yet th i s particular spot seems to be her cho ice and
the re i s someth ing about it w hereby she can remember i t
and distingui sh i t from all the rema inder o f the large area
in return i ng to i t again and again H ere s he at once begins
her digg i ng In this task s he always works w ith an i ntens ity
which makes he r appear to b e driven by some fury S he
cannot wait to calmly ca rry out load a fter load o f d i rt as
her S i ste r sp ecies do but a fter hav ing to rn the earth loose
w ith her mandibl es s he scratches or kicks it back w ith her
forelegs straight between her h i nd legs and far out beh ind
her i n a spurting stream o r j et Much o f the time when
she i s working on o r near th e sur face o f the ground s he
throws an almost unbroken stream o f d i rt out behind her
as s he digs o ften hurl i n g i t a s fa r as ten i nches back o f
her The front l egs are prov ided w ith bristles which serve
as brushes o r rakes w ith wh i ch she sweep s back the di rt
T o do th is s he i s compelled to stoop forward ; thi s g i ves
her a comical appearance o f sweeping w ith her elbows

as the P eckhams suggest for anothe r species o f B em
O ften S he seems dissati sed w ith he r location and makes
several beginn ings i n as many d i fferent places although we
have never b een abl e to see the sl ightest reason why the
.
20
was begun on July 1 4 1 9 1 5 When w e excav ated it on

July 2 6 twelve days later it contained a larva three or four
days old togethe r w ith four ies
S ince the cell at the
bottom must have been completed be fore the egg was de
pos ited thi s l eaves eight o r n ine days fo r the construction
o f the nest and the incubation o f the egg Thi s seems in
deed a long time for so rap i d a worke r as B n ubi lip en n is
We have occasional ly found her resting or loite ring in he r
incomplete o r newly n i shed burrow and we k now that

s he doe s not wo rk on cloudy or rainy days ; so w hen we
have made ampl e allowance o f time for these indulgences
thi s may a fter al l be a fai r estimate o f the normal time
devoted to the making o f a nest
The no rmal burrow th ree eighths inch i n diameter at
the top p ierces the ground obl iquely and cont i nues down
ward at an angle o f about 30 to 4 5 w ith the sur face
fo r ve o r s ix inches to its h orizontal oval terminal pocket
wh i ch i s one inch long by one hal f inch i n d iameter Thi s

cell i s holl o wed out more f rom the roo f than fro m the bot
tom o f the burrow The tunnel s a re usually approximately
stra ight and only occas ionally swerve to the right or the
le ft ; the slope o f the tunnel rarely vari es
Most o f the nests are normal as the rst two illustrations
in g 3 but there a re occasional exceptions
O ne nest
went down at an angle o f
the tunnel was about twelve
inches long wi th the usual pocket one i nch by one hal f
inch ; anothe r nest had a normal gallery but a cell an inch
and a quarter long and one i nch w i de Another was normal
at the entrance but soon curved straight downward and
B ut
then back unde r itsel f ( lower illustration i n g
the most novel form o f nest which w e have yet d iscove red
was one which started down and eastward at an angle o f
fo r two inches ; then turned sharply to the south and
downward at an angl e o f
for four inches ; then con
.
'
TH E
FI G
T he b u r r o
t i n ued
s of
B E M B I CE N E
em b ix
n u b i li
WA S P S
en n i s
T w o -thi rd s
21
n a tu r a l s i
ze
south at an angle o f
fo r tw o inches to the un
ni shed pocket where the owne r was at work It is strange
that she shoul d stil l be extending it a fter s he had worked
on it for seve ral days and it w a s al ready beyond normal
depth
While w e have opened many nests during the two years
and f ound them in vari o us conditions and o f sl ightly varied
,
22
forms w e have never found m o re than one cha mbe r at the

bottom o f the tunnel and one immature wasp there in The
only ne st having more than one organism w a s one which
contained a medium sized larva feeding on ie s and a much
smaller second one i n the same and only chamber The
sl ight resembl a nce o f the two larvae caused us to doubt
whether they were o f the sam e spec i es although the addi
O n another o c
t i o n al one w a s a Hymenopterous larva
cas ion in d igging up a burrow w e found a second one in
the earth close bes ide the chamb er that conta ined a large
larva ready to sp in but thi s p robably di d not bel o ng to the
same nest The wasp s i n the i r sem i gregar i ousnes s o ften
build the i r burrows i n close j uxtaposition s o the cells o f
d i fferent nests seem to have only a th i n partition o f earth
between them
3
P arke r however nds that h i s B n ub i lip en n is has hab i ts
simila r to P h ila nthus fo r he says : The burrows

e nte r
the ground at an angl e o f
At a d is tance o f from e ight
to twelve inches from the entrance lateral branches are g i ven
At the
o ff w hich serve a s b rood chambers for the larvae
time o f my observation no burrow w a s found w ith m ore
than ve o f these chambers ; most had four and a few had
only three I n the chambers more than one larva may be
reared i n wh i ch case the rst i s placed at the extreme end
o f the chambe r and when full grown and encas ed a wall
is placed across the chamber and anothe r larva reared b e
tween th is and the main part o f the burrow
These
observation s were made on August 1 8 and 1 9 and although
many burrows were opened only l a rvae we re found Many
,
Ohi o N a t
s p ec i e s
in
1 0 : 1 63 1 6 5
-
1 8 72 ,
l o ic a l d a t a
on
t hi s i s th e
t hi s
D e sp it e the
o n ly
s p ec i es ,
S t Lo ui s 6 : 4 5 8 1 89 4 w ho
o f f o ur s p ec i e s o f p l a n t s
.
f ac t
that
p ap e r t ha t w e
x c ep ti n g
f o un d
th e
'
C re s s o n
nd
c an
R o b ert s on
w hi
ch
ha s b i o
T ra n s Ac a d S c i
t f eedin g
a d ul s
d es c ri b ed the
on
th e o
ers
WA S P S
23
o f these had completed the i r growth and were encased in

cells
but in every instance an immature larva in some
stage o f development w a s also found in the burrow In
no case however w a s more than one develop ing larva found
i n any burr o w I n one burrow with four branche s three
contained matured and encased larva e and the fourth
j ust newly constructed conta ined tw o recently killed
house i es on one o f which was found an unhatched egg
From the data given above it would appea r that the wasp
rears o nly one larva at a time

Thi s interesting note recorded f rom W ilson Kansas
which d i ffers s o w idely from our observations on the num
ber o f chambers to each burrow shows that the wasps o f
D r P arker s l ittle colony o f n ubi lip en n is have acquired the
economical adaptation o f us ing t hei r bu rrows fo r more than
one chambe r It has been a matte r o f doubt whethe r B em
b ix make s one burro w and carries that larva through i ts
in fancy be fore beginn ing the next or wheth er she keeps up
mo re than one domicile at a time The P eckhams think
from s ome expe riments that B s p i n o la e takes care o f only
one nest at a time and from our observations on the per
manent closing o f the nest early i n the season w ith only a
single young o n e withi n we think i t probable that i n our
colony the second burrow i s constructed only a fter the rst
has been completed Whe n w e cons ider ( as will appear
later ) how poor the dissemination I S in thi s species we can
easily s ee how a digressio n o f making three to ve chambers
to a tunnel may eas ily become a habit and nally a family
characteristic and may become constant i n the colony D r
P ar k e r states that these wasps had nest ed in this spot a n
n ua l ly fo r a number o f years
H ence the establ i shment o f a very p retty and e c onom ical
hab it At the same time one must recogniz e a n ew hab it
correlative o f the multipl icity o f cells having arisen he re
.
W A S P ST U DI E S AF I ELD
24
which l ike the new habit j ust described woul d go enti rely
outs ide the genus B em b ix We know that the digger wasps
B emb ix A m m oph i la P ri on on yx are essentially diggers
but D r P arker nds the B emb ix in his colony taking up
mason work also building partition s to form new cells
A wall i s placed acro ss the chamber and another larva
reared between this and the main part o f the burrow

It
seems incredible that n ub i lip en n is coul d build the wal l o f
dry sand or dust unless s he too has acqui red the water
carrying habits o f certa in O dy n erus The ll ings o f loose
dusty earth such as sh e scoops into he r holes he re coul d
never be made to serve as a wall across the bulging m iddle
o f the chamber I t w ould be well worth wh il e to make an
e ff o rt to s ee how general th i s new hab it has become
We have s o many times found the mother B emb ix in her
burrow when w e opened i t at any time near or a fter sunset
that w e have arr ived at the conclusion that s he sleep s in
h er nest when s he has one We have al s o found her here
safely hidden away on dark o r rainy days fo r B emb ix
loves only warmth and sunl ight The tunnel i s always tem
r
l
r
i
closed
under
these
conditions
Thi
s
i
s
accompl
ished
o
a
p
y
by the B emb ix go ing down into the hole and push ing up
loose di rt from below until the mouth o f the burrow i s com
l
l
t
closed
One
mo
rn
ing
we
found
that
the
dashing
rai
n
e
e
p
y
during the n ight had packed the so il hard and washed down
the loose earth leaving the hol e open We arrived j ust i n
time to s ee the mother B emb ix cutting a small area about
the open ing and kicking the d irt thus loosened int o her bur
row until the aperture was closed
I t seemed that th is
mother had remained in her burrow all n ight w ith only the
usual temporary covering over i t and that the rain had
washed down th i s dust covering and s he unwilling to have
hersel f and her nest exposed thus to danger was hastening
to cove r her burrow again even though s he had to dig up
,
'
TH E
B E M B I CE N E
WA S P S
25
the hard packed earth w ith he r j aw s to o btain the meces

sary l ling
The same morning another B emb ix al ighted upon a par
t i c ula r spot wh ich to us was absolutely indistinguishable
from all the rest o f the smo o th washed d iamond

S he fell
industriously to digging and in a moment opened her tun
nel and entered S he had evi dently been away from home
during the sto rm and found shelter elsewhere Even this
new condition o f the soil about the nest di d not con fuse
her S ome m inutes later upon returning w e we re surp ri sed
to see the hol e aga in covered and the topography smooth
but about s ix inches f rom the hole ( the place was marked
o r it would have been quite indistinguishabl e ) was much
loose di rt which had been swept well away fro m her door
On one occasion o f exceptionally heavy rains the earth from
the upper side o f the area washed down and deeply covered
every trace o f thei r nests ; but be fore evening about a dozen
had worked the i r way out o f the i r holes none the worse
fo r wet weather
I t was on J uly 2 o f the rst year o f our observations
or j ust t w o weeks a fter the rst emergence and danc e o f
B emb ix that we dug out the rst nest that was com p leted
and occup ied The waspl in g was only a tiny larva one
fourth inch long and w a s f eeding upon its rst y a del i
cate l ittle yellow w inged D ipteron
On many occasions
the rea fte r when w e found a very tiny larva feeding the
prey invariably w a s a very so ft del icate l ittle yellow or
green w inged y not the coarse heavy kind such as house

ies stable ies or horse i es which the mothers bring in
when the larva i s stronger
Can human intelligence do
?
a
better in del icate m ternal care
It was startl ing at rst
to s ee a tiny larva and y fai rly rattl ing a round i n so

large a cavity ; but i t only p roves again the maternal sol ici
tude o f M other B emb ix who it seems knows how to est i
-
26
mate the future needs o f space fo r her growin g in fant its

food and the usual res idue
We have already stated that this wasp stays w ith her
nest and daily brings in enough f resh ies to mee t the
needs o f the young one until i t reaches maturity and p u
pates Just how many ie s are brought in each day w e
have neve r ascertained w ith accuracy but we suspect that
the food supply varies w ith the needs o f the i n fant Also
as w e shall s ee later th is instinct o f the mothe r for pro
v i d i n g the daily food for her young i s sometimes de fective
and leads the mother to d o various unprotabl e things
Howeve r what s he does i s s o wonde r ful that w e shall not
p resume to cr i ticise her fo r occas ional fa ilures or erratic
behavio r
When forag i ng and bri ngin g i n ies they come and go
w ith surpri s ing rap id ity o ften ma king three or four trips
i n twenty m i nutes They al ight upon the nest w ith the y
clasped between the
s o well concealed unde r the i r bod ies
m i ddle legs that w e can hardly s ee it They neve r waste
a second s time i n searching to and fro for thei r nests but
drop down f rom ight di rectly upon them every t i me Thi s
homing instinct i s truly marvellous w hen we co nside r that
the bare area occupied by the i r colony i s devo id o f any ap
parent landmarks by wh ich they could locate the ir ho les
ami d the numerous hole s o f othe r insects They wo rk fast
and furiously f rom the i nstant they arrive probably to
evade paras ites w h i ch soon shadow them i f they are more
than a moment in thei r work or to escap e attack from thei r
s isters which are hot a fter them I n a fraction o f a m inute
they d i g through the temporary closure o f the burr o w and
scrambl e i n w ith the y ; in a f ew s ec o n ds they emerge
head rst turn around at the brink o f the hole and re
enter at once
Fo r an i nstant thi s p e r formance seem s
strange but only fo r an instant ; they enter j ust far enough
,
28
food for thei r young I n contrast to thi s we see the nal or

permanent closu re wh ich i s made only at the end o f the
la rva s feeding period The contrast i n the method s o f e f
f ec t in g the two closings w ill be at once apparent
When the wasp has n ished feeding the larva she eme rge s
f rom the open hole but does not turn around and go right
back in head rst as p reviously I nstead o f going in and
kicking the di rt up from the lower part o f the channel she
begins scratching and kicking the l o o se d irt from the s ur
face into the hole behind he r moving forward as she does
.
'
a)
FI G
em b ix
n u b ili
f o releg s s w e ep in g the
N a tu ra l s i z e
a im
.
lli g
e n n is
s o il
her b u rro
b o dy
un d e r t he
and
N o te
t he b r u s h- l i
th e a c c u ra c y
of
h er
so until she i s two or three inches away from i t Then s he

runs to the hol e peeps in turns around and backs in kick
ing scratching clawing and crowding the di r t deeper down
into the hole and out o f s ight Then s he comes out and
exactly repeats the whol e per formance several times each
time beginning at the margin o f the burrow to scratch the
so i l back in and wo rking gradually outward w i th her face
h
s
away from the opening
Fig
Thus
continues
e
(
until the hol e i s lled s o near the top that she cannot back
.
WA S P S
29
down into it any more ; then she assumes the same pos ition
backwards on the hole and crams and packs the di rt down
w ith her hind pai rs o f legs and pounds rubs and punches
i t down with her abdomen ; then brushes more di rt back
upon the depression and repeats the p acking process two o r
three times until the ll i s brought up exactly l evel w ith
the sur face o f the ground Having nished tamp ing the
earth s he begins sweep ing al l around as usual to remove
all traces o f he r nest from the sur face o f the ground H e r
conduct at thi s point varies with the conditions ; i f there
i s a surplus o f loose dirt near the s ite o f the hole s he
sweep s i t well away and scatters it evenly over a cons ider
able a rea ; i f i t i s bal d s o that traces o f the lled hole may
be seen s he sweeps other dust back ove r i t from a d istance
unti l no trace o f a scar may be seen When sweep ing away
f rom the hole she begins at the oute r edge o f the patch o f
di rt faces the hole and k icks the dust back o f he r as s he
moves from s ide to side and gradually nears the opening all
the time with a stream o f dirt spurting out beh ind her ; then
she backs o ff to the periphery o f the dusty space again b e
fore beginn ing anew scattering it furthe r and furthe r from
the nest I f however s he i s sweep ing di rt toward the hole
she di splays far greate r skil l in estimating the distance and
the force necessary to throw the di rt in precisely the right
spot She always works w ith the hole squarely behind he r
so she can throw the d i rt back o f her She works fast and
furiously in the hot s un brushing lightly when near the hole
and in c rea s m g the strength o f her throw as s he moves
further away f rom i t and w ith wonder ful j udgment o f
di rection and di stance o f throw h itting the region o f the
burrow w ith astoni shing accuracy f rom various di stances
a s she works
S
ee
g
She
move
s
f
rom
s
ide
to
s
ide
(
so that her path p ractically describes arcs o f ever widen ing
circles the center o f which i s the burrow When she has
,
o
3
reached the l i m it o f her di stance f rom the hol e she does

not wander aimlessly over the area but always returns to
the hol e to begin a new series o f arcs Th is marvellous
system o f cover i ng the ground i s more eas ily made clear
by the accompanying diagram ( g
I t i s interesting
to note that the wasp usually con nes i ts sweep ing to the
area in front and at the s ides o f the hol e but i s rarely seen
working beh ind the hole O f course th is may be due largely
to the fact that the loose dirt all l ies in front o f the burrow
where i t has be en th rown dur i ng the digging
Thus s he sweeps spreads and red i stributes the loose s ur
face dust for a foot o r more until i f we have not the exact
location o f th e nest marked we can seldom nd it so well
i s it concealed Then s he c i rcles t o and fro on the W i ng
an inch o r s o above the area survey i ng it care fully fo r s ev
e ral seconds and goe s whi rl ing o ff i rtat io usly w ith the
o ther wasps
wh i ch f rom t ime to time have been dabbing
down beside her and bump ing her One B n ub i lip en n is
accompli shed the permanent clos ing o f her ne st i n twenty
m inutes but they usually requi re a longer time to ni sh thi s
critical p iece o f wo rk
When w e s ee such highly developed instinct for the care
and sa feguarding o f the young w e are surpri sed to nd
that thes e mothers are o ften guilty o f comm itting gross
e rrors i n the most fundamental points We watched one
wasp working w i th great care and precision i n clos ing
he r nest
A fte r the last supe rcial trace o f the hole had
been care fully obl iterated and the surrounding ground swept
clean and s he had departed w e op ened the burrow in the
full expectation o f obtaining a mature larva but we were
shocked to nd that the larva had pupated long ago and
had been dead and rotten for some days at least No fresh
or uneaten ies were i n the chamber ; only the old d bri s
What s he could have been doing there o r why she was so
,
B E M B I CE N E
TH E
WA S P S
T
3
:
I
v\
x
!
l
"
@V
I
l
I
I
I,
I
l
l/
l
I
[
I
r
FI G
i p en n i s i n
c l o s ure
D i a g ra m
s
eep i n g
s ho
th e
wi
ng
th e
d us t b a c
s y s te m a ti c
I
.
m e thod
i n to h e r b urro
of
B em b i a
'
n ub i l
f o r the p erm a n en t
2
3
parti cular in seal ing up that dead larva w ill always be a

mystery Another mother spent equal pain s i n permanently
clos ing a burrow wh ich late r p roved to be stark empty In
yet another seale d b urrow w e found bes ides the newly
pupated larva a spoon ful or mo re o f d ebris w ings and
legs and besi des thi s s ixteen whole untouched ies show
i ng how her sol icitude had oversuppl ied the larva The ies
were al l dead ; never i n deal ing w ith B n ub i lip en n is di d w e
nd them al ive or paralyzed nor w ith the thorax crushed
In th i s nest w e found also a pai r o f w ings o f the bee y
S p og os tyl um a n ale S ay wh ich i s paras itic on mud wa sps
but here i t had been brought in fo r food
The contents o f the chambe rs and the occupancy o f the
holes by the diggers are l ittle understood ; as yet the l ine
o f di fference between the no rmal and the abnormal use o f
the burrows ha s not been drawn
N ot a few hol es have been followed to the i r end only to
nd the nest all ready for occupancy and the mother wasp
calmly resting at th e bottom Unde r these circumstances
the hole wa s usually cl osed fai rly rmly from w ithin
O f course i t may be that w e chanced to i nterrupt he r
at a time when s he had j ust completed he r burrow but
thi s condition occurred s o frequently that such a chance
woul d have been h ighly imp robabl e ; hence w e suspect that
to
s he uses he r burr o w not only for n i d i c a t i o n but also
a cons i derabl e extent a s he r o w n dom icile To be sure
the gallery near the mouth o f the burrow i s always closed
by a mound o f loose dirt when the wasp i s digging the lowe r
part o f the hole but then the activity can be detected at
short i ntervals when the d i rt move s or humps up from the
-
Ou
i t w a s i m p o s s ib l e t o
n e st
lo n
r ev i s i t
o
gh t
en o u
m ar
ed
n ests,
b l i te ra ted them
f req uen t
th e
of
a c c o un t
eep
g et
in
us e
of
c o n n ec ti o n
th e
w i th
c o n n e c t e d s t o ry
b ut i n
ev ery
case
el d
on e
E v ery
th e b o y s
b y th e b a l l- p la y er s ,
m o th er
e ff o rt
w i th
t he
or
h er
on e
w a s m a d e to
sp
ed
s ho es
TH E
B E M B I CE N E
WA S P S
33
pressure o f the wasp pushing up from beneath or occasion

ally the wasp itsel f wil l come p ush ing its w a y out backwards
through the loose so il emerging not a l ittle dishevelled and
shake itsel f vigo rously to free it from the cl inging d irt pre
c i s ely as a dog shakes h imsel f upon coming out o f hi s bath
S he then p roceeds to dig out and sweep back the dirt she has
brought to the top o f the hole B ut at these resting periods

mentioned above no such movements are seen f or days
at a time s o we suspect that s he i s merely resting and
B ut i s s he only
enj oying the security o f her home
grati fying a wh im or may she be waiting for the matura
?
tion o f he r ova o r some other physiological phenomen o n
The population dim in ishes at the end o f the season ; perhaps
the old adults s imply go o ff and die a fter they have com
l
i
i
i
o
n
e
t
d
thei
r
seas
on
o
f
n
d
c
a
t
e
p
On this occasion w e might mention one other form o f
mor tal ity which i s yet unexplained Early each year when
digging out the burrows o f various species w e come u p on
a few subterranean chambers which have never been
opened containing a dead B emb ix Whether these are
y o ung wasps wh i ch have sa fely arrived at mat ur i ty but have
died be fore eme rging as o ften happens i n other species o r
whethe r they are parent wasps which have died while in
the burrows w e had no way o f ascertaining
The contents o f a few nests have puzzled us utterly One
hole had long roused ou r curiosity as we watched and
waited for i ts permanent seal ing Finally one evening at
twil ight w e broke open its temporary closu re and found
therein six B n ub i lip en n is ready to spend the night Two
o f these were captured and p roved to be females S ince we
now know o f the male p riority o f emergence and d isappear
ance w e feel fai rly sa fe in assuming that all o f these were
,
T he S l e ep
of
I n s ec ts
An n E n t S o c Am er g : 2 40
.
916
34
females On one occasion we s aw a B emb ix hav ing much

di fculty in keep i ng out a si ste r wasp whose home it seems
had been destroyed and who did not seem to care that it
was not her own hole which s he w a s entering Th is gives
us a poss ible clue to an explanation o f the presence o f s ix
wasp s in one burrow ; p erhap s they had returned to the el d
at tw il ight found themselves homeless a fter the ball game
and had found i t conveni ent to crowd into the same open ing
for a n ight s shelter
The B emb ix do not seem to b e so discriminating in re
ga rd to the choice o f thei r p rey as are som e oth er wasps
S o long as they can p ick up ies they do n o t s eem to be
particula r about the variety We have f ound i n their bur
row s : L uc i li a c a esar L i nn [ F Knab ] S p og os ty lum ana le
Say [ E Knab ] S a rcoph ag a s p [ F Knab ] S p a rn op oli us
b revi r os tris Macq [ F Knab ] house ies stable ies and
many more species wh ich coul d not convenient ly be
identi ed
The amount o f food consumed by one o f these youth ful
gourmands i s quite astonishing One wh ich we brought up
from i n fancy by han d consumed seventy n ine house i es

be fo re it spun its cocoon Another consumed the forty n ine
house i es and one large stabl e y which had been given it
and then attempted to sp in i ts cocoon In the smooth tin
A fter that
box i t made only a silky carpet ( g 6 a )
th irty s ix more house i es were intr o duced and i t devoured
twenty s ix o f them in sp ite o f the fact that i t had already
begun to spin H ow many ies the mother p rovides for a
single larva has never been ascertained ; but a fter its pupa
t ion there i s usually f rom a teaspoon ful to a tablespoon ful
o f legs and w ings remaining O ccasionally we nd that
a mother has devotedly b rought in mo re ies than the larva
has been abl e to consume ; but some o f our wo rk l eads us
to suspect that these gluttonous young are more o ften w ill
.
6
3
Our present evidence indicates that there i s Only one gen

e ra t i o n pe r year o f B
n u b i li p en n i s
From the time o f
emergence in J une or early July the females go o n w ith
n i d i c a t io n until the m iddle o f S eptember
We do not know
how many nests each constructs during this sea son but they
ought to make several i n orde r to o ff set the high mortal ity
and keep up s o large a population The numbers gradually
d imini sh as autumn approaches ; s ince w e never s ee them
elsewhere hereabouts w e conclude that the mothers grad
They spend the w inter in the pupal stage and
ua lly die o ff
desp ite the fact that they reach th i s stage at di ff erent dates
they all emerge s imultaneously when the warm rays call
The persistence with which B emb ix adhere to the one
location wh ich w e have watched year a fter year is aston is h
i ng Truly the i r d is sem ination seems to be n i l In regard
to th is species at least one i s soon ready to agree w ith the
P eckhams in thei r op in i on that wasps general ly do not travel
far f rom the place o f the i r b irth H ere the wasps o f o ur
parti cular colony have had at thei r d isposal a pasture equal
in s ize to s ix ci ty bl ocks i n wh ich to choose the i r nesting
s ite yet yea r a fter yea r they l im it themselves strictly to two
di stinct corners o f a base bal l diamond in the mi ddle o f
this eld H ere they su ff e r weekly and sometimes daily
persecutions unde r the sp iked shoes o f the boys ; they are
routed f rom the ir work i n the m iddle o f the day an d thei r
nests are obl iterated in the powde red earth and ri ot a few
must su ff er death in the m el ee ; yet not even this pressure
causes them to move twenty ve feet to one side wher e
they m ight enj oy peace and p ro sperity The fact that each
success ive generation shoul d S how the same cho ice and the
same p ers istence i s all the mo re astoni shing I n another
vacant lot a m il e away was another colony which to our
knowledge pe rsisted in l ike manner on a small area for
three years Finally a building was er ected on the site and
.
TH E
B E M B I CE N E
WA S P S
37
the B emb ix population must have been exterminated for

a fterward the count ry roundabout was scou red i n search
o f them but no trace o f the ir hav ing settled el sewhere was
ever found although j ust across the road from the lot was
a large tract o f vacant land thoroughly suitable to the i r
needs In the tw o colonies in co rners o f the baseball dia
mond the numbers are becom ing fewer and fewer so i f
they do not mi grate they w ill p robably s o on meet w ith ex
te rmination
,
B emb ix s p i n ola e Lep [ S A Rohwer ]

.
Othe r observers notably the P eckha ms and P arke r have

already wri tten upon the li fe o f thi s species so we o ffer no
new data excepting a contributi o n o f one or two eccentric
circumstances
One burrow which we di scovered on Howard s H ill on
S eptembe r 2 5 1 9 1 6 seemed externally a typ ical B emb ix
hole I t pi erced the earth diagonally and much loose sand
was strewn about the mouth o f the burrow It went d o wn
thus for an inch and a hal f then turned abruptly at right
angles and from there i t c o ntinued downward in a spi ral
so that the pocket w a s di rectly beneath the entrance The
total length o f th is sp i ral burrow w a s S ix inches and the
term inus wa s three inches below the sur face We sa w no
obstructions in the s oil wh ich w o uld necess itate such con
tortious in the channel I n the no rmal nest ( acco rding to
P eckhams ) the tunnel goes down obl iquely to a po int from
three to ve inches below the sur face o f the ground
At the bottom o f the hole we found the mothe r B emb ix
togethe r w ith a med ium sized healthy larva thirty large
7
B ul l Wi s c
P ro c U S
.
G eo l
N at
N a t H i st
.
M us 5 2
.
S u rv
: 1 2 7- 1
31
2 :
5 8 72
91 7
1 89 8
WA S P
8
3
S TU D I E S A F I EL D
l iving Diptera larvae four adult bee ies S p arn op olis

b r evi r os tri s Macq [ F Knab ] the remains o f e ighty v e
more o f the same and the remains o f one L ucilia c a es ar L
Judging f rom the enormous amount o f food
[ F Knab ]
rema in s when the feeding stage o f the larva w a s as yet far
f rom completed and the p resence o f a healthy medium
s i zed B emb ix w e can only conclude that the Diptera were
no t true paras ites s ince they had not attacked the larva o f
the B emb ix but me rely inquil ines o r imposto rs w ho depen d
up on the mothe r B emb ix to work extraordinarily h a rd to
keep them as well as her own young suppl ied w ith meat
What a terrible toll that s he must p rov ide constantly for
thi rty others in order that he r own may l ive !
One wonde rs w hy the D ipterous larvae so many in num
ber spare d the B emb ix larva fo r it certainly would have
made a ne meal We do not w i sh to speculate excess ively
yet i t i s probably only fai r to breathe ou r susp icions on the
subj ect I f they shoul d devour the B emb ix l arva it would
satis fy them fo r only a short tim e but w ith the dem ise o f
the mothe r s child the i r further supply o f food would b e
cut o ff since p robably the right ful in fant i n the cell i s the
stimulus for the mother to continue replen i sh ing the food
supply for al l o f them as fast as i t i s exhausted With
these thi rty one mouths to feed da ily i t i s l ittle wonder that
the mothe r had to sacri ce all o f her lei sure that might have
been appl ied to imp roved methods o f domesti c economy
We d o not p retend to s a y that the y larvae knew the
resul t o f thi s action and behaved accordingly and numerous
othe r i nstances o f the same kind woul d be requi red even to
p rove that thi s i s a case o f habit o r bl ind instinct ; yet i f i t
were proved to be such i t would be no more wonder ful than
many o f the del icate adaptations al re a dy known to exi st in
the i nsect world
-
WA S P S
T H E B E M B I CE NE
Mi cro b em bix
39
mo n odo n ta Say [ 8 A Rohwer ]

.
We s a w thi s l ittle banded M i c r o b em b ix open her hole and

ente r and close her nest preparatory to leaving whe reupon
we captured her The hole went down diagonally for seven
i nches then turned at right angles and continued its down
ward course for s ix inches more where it term i nated w ith
o ut e ithe r pocket o r prey
The end was s ix and one hal f
inches below the sur face o f the earth
A second one was seen to l eave her hole a fte r care ful ly
covering i t up The illustration ( g 7 ) tells better than
words the nature o f the burrow The nesting s ite w as the
sem i barren sandy area ( g 8 ) along the river at Lake
V iew Kansas The total length o f the hole was fourteen
inches The pocket w a s hollowed out from the ceil ing o f
the burrow G reat was our surp rise upon nding in here
two beetles N eoha rm o n i a veluti n a [ H S B arber ] and
H ipp od a m ia I 3 punc t a t a [ H S B arbe r ] two tree hoppers
D ra ec ulac ep ha la m olli p es Say [ E H G ibson ] and P hlep s i us
i rr or a tus Say [ E H G ibson ] one sp ide r a Xys ti c us ner
L S hoemaker ] and o n e ant P he i dole
v os us B anks [ C
All s ix items o f prey
vi n ela n di c a Fo rel [ W M Wheele r ]
were dead dried and hard No egg was found w ith the
food We can hardly indeed call this c o llection by the
term p rey because it was only a collection o f dead carcasses
which the wasp had garnered H artman too nds that M
mo n o do n t a takes home i nsects that are already dead H is
l i st includes : slender red caterp illars the l eg o f a grasshop
per small queen ants large red ant ies o f various kinds
bugs belonging to ve di ff erent genera tree hoppers poli stes

wasp f reshly killed grasshopper a dry Muti ll id and old
O rthopterous pupal case s with dry dead pupae i nside A
lengthy and very interesting account o f the mating behavior
the nesting habits food habits etc o f thi s species i s included
,
WAS P S TUDI E S AFI ELD
4o
FI G
T he b urr o
of
M i c r o b em b i x
m o n o d o n ta
in the paper by P arker

H i s records o f nest p rovi sioning
based on several observations di ffer f rom ours above H e
8
P roc
N at M us 5 2
.
: 1
34
1 40
91 7
T H E B E M B I CI N E
WA S P S
41
says that when the burr o w i s completed a s ingle egg I S placed

in the brood chamber at the extreme end Th i s i s rmly fast
ened in an upright position in the sand o f the oor No
food i s placed in the burro w until a fter the egg has hatched
in the cou rse o f two or three days O ur S ingle wasp had
al ready p rovi sioned the nest w ith s ix dead insects and had
le ft the nest p resumably for more but a very care ful search
revealed no egg nor larva Whether the Ohio wasp s di ffe r
in the i r ways o f housekeeping from the Kansas wasps o r
W hether ours w a s only an eccentric ind ividual we do not
know The crooked nesting burrow described above seems
also to be only an eccentricity These points only S how that
wasps as well as other beings may express consciously or
unconsciously the 1 r i ndividual ity
B arth
in speaking o f this genus and al so o f B emb ix
says the Larvae are fe d from day to day nests remaining
open during excurs ions

Th is sweeping statement cann o t
apply to M i c ro b em b ix mo n odo n ta
,
B i cy r tes q ua drif as c i a ta
Say [ 8 A Rohwe r ]
.
In a sem i barren sandy area ( g 8 ) beside the Kaw

River at Lake V i ew Kansas we found our only specimens
o f B q ua drif as ci a ta P arke r ha s described very well her
manne r o f app ro ach ing her nest as she returns to it S he
returns on the w ing ying h igh in the ai r and poi ses in m id
ai r ten or twelve feet above the hole then d r o ps straight
down as i f w ith a parachute digs a moment at her very
feet where S he al ights and 10 ! her burrow opens Truly
the students o f animal psychology w ho are trying to solve
the hom ing o f insects have here an inte resting problem
-
1 0
B ull Wi s c
.
10
Loc
c it
N at
1
34
H ist
S oc
: 1 1
910
WA S P
2
4
S TU D I E S A F I EL D
She f ollows the same method in departing f rom the nest

or i n starting homeward from the eld when foraging She
ri ses straight up in the ai r s o h igh that it i s di fcult to fol
low her with the eye i n the dazzl ing sunlight then darts away
ho ri zon tally above the heads o f most living things
We once watched one climb ing in and out among the
branches o f a cocklebur plant carrying her l ittl e bug snugly
beneath her resting once o r tw ice on a lea f All at once
w ithout p rovocation s he leaped up into the a 1 r and ew
straight upward highe r higher unti l we lo st her in the
dazzl ing sunl ight but in a few seconds she came down from
her highway i n the skies straight down to a spot only ten
feet distant from where s he had started Here s he scratched
on the ground at h er feet opened her hole and w ent in w ith
out letting go her p rey S he remained w ithi n for about
fteen m inutes and closed the hole l ightly behind he r w ith
loose sand when s he came o ut be fore departing upward
The prey whi ch she brought into the nest was in all
cases observed a smal l bug S he clasped it beneath her
abdomen and in the one or two cases wh ich we have been
able to observe m inutely carried it w ith the bug s ventral
s ide toward her o w n The four bugs found i n one nest
we re all somewhat S imilar but o f d i fferent speci es all
nymphs o f P entatomidae [ E H G ibson ]
The two holes which we dug out w ere unl ike They re
sembled a B emb ix n ubi lip en n is hol e in general appearance
penetrating the ground at an angle o f thi rty to forty de
grees w ith the sur face One was twelve inches long and
straight ; the other was seventeen inches i n total len gth and
the large terminal pocket was n ine and a hal f inches bel ow
the sur face o f the ground The burrow curved i rregularly
forming hal f a spi ral
.
W A S P S TUDI E S AFI ELD
44
Sphec i us sp ec i os us
1 1
Drury [ S A Rohwer ]
.
Th ree individuals o f this species in company w ith D ip

tera and a P ap i li o buttery were bus ily feeding upon a mass
o f foamy s a p that exuded from a wound near the base o f a
huge s un o w er stalk They were determ ined to get the
s ap
and when we attempted to take them they expressed
thei r indignation at be ing disturbed by the loude st no is e that
w e have ever heard a wasp make They buzzed and ew
about and always ma de attempts to return to thei r b ac
chanal The s a p seemed to have fermented in the summe r
heat s o perhap s l ike ensi lage i t was particularly exciting
to an imal a ppetites
The next day w e again found these 5 s p ec i os us attracted
to the s a p greedily drink ing thei r ll a n d b o i s tero us ly buzz
i ng bump ing and whi rl ing around at i nterval s
N o w they
had fo r thei r compan ions several horse ie s Ta b an us exul
O S [ F Knab ] many smaller D iptera one cl ick beetle

Lud i us a tten ua tus [ J A Hyslop ] a hal f dozen P ol ist es
a n n ul a ris one spotted cucumber beetle and one blue P ap i l i o
H enc e i t seems that the i r pulque w a s popular 1 n i nsect
society
H unger ford and Will iams found that th i s wasp quar
rel ed w ith C hl o ri on c a erule um fo r the sap ooz ing from the
w illows Riley has worked up a beauti ful account o f the
behavio r o f th is speci es a popular version o f w h i ch may
be f ound i n H oward s Insect B ook pp 2 2 2 5 or Weed s
Li fe H i stories o f American Insects pp 1 5 7 1 60
,
'
12
1 3
1 1
1 2
1 3
0 f th e
En t
s ub -
N ew s
I n s ec t Li f e
f am ily
2 3 : 246
S ti z in a e
.
19 12
: 248-2 5 2
89 2
CHA P TE R I I
B EHA VI OR
O F W ASPS B E LO N GI N G
P O M P I L I D AE
P o m p i lo i d es trop i c us
THE
To
Linn [ S A Rohwer ]
.
F A M I L!
One m ild sunny S eptember mo rn ing w e were walking

acros s the eld w hen we were startled to espy a telltale hole
i n the path I t w a s only the beginning o f a burrow but
i n a mo st hazardous s ituation r ight i n the beaten path ove r
which hundreds o f workmen passed da ily and also at its
intersection w ith the wagon road yet because the spot was
vo id o f vegetation and sunny the wasp had proceeded to
make her nest there as i f unaware o f any danger The
ground was dry and hard enough almost to break a pocket
kn i fe to dig i n it while the sur face had been ground b y
the t read o f many feet i nto deep ne dust The hole w a s
perhap s one hal f inch deep ; the sp ider occas ionally twitch
ing lay only an inch away From this it i s at once apparent
that thi s wasp does not dig the hol e rst and then fetch he r
p rey fo r th i s hole w a s only begun and the spide r an im
mature mal e L y cosa f r on di c ola Em [ N B anks ] lay in
front o f it
The black P omp il id conspicuous against the grey dust
w ith its broad orange band on the abdomen w a s seen
hunting d iligently all about the v icinity eight o r ten feet
away She returned to the hole once but we had tampe red
with it knocking in a lot o f loose dust in trying to break
through the hard crust o f earth fo r her s o she le ft it at
,
45
46
once and continued her search s ix feet to the west She

eventually came back and tried to dig i n three or four places
near her original locati on but the loose sur face d irt always
rol led in upon her She le ft again and wandered on foot
down the road ; nally a fte r ten m inutes she returned on
the w ing found the spot w ithout d i fculty and again dug
i n a hol e wh ich p romi sed to be success ful A caterp illar
hurried by very near to her and seemed to make he r uneasy
s o s he drew he r sp ider back three o r four i nches
Again
M ean
s he departed fo r a wh i le on an unexplained j aunt
while w e brushed the loose sur face dust aside to give her
access to the soil which w a s rm en ough for he r to dig
in and w ith a kni fe began a new hol e H e r S pider still
lay near by A parasiti c bee y wh ich had been hover
i ng about inqui s itively from time to time h ove red over our
hol e the s ame as over hers The wasp cam e back thr i ce but
d i d not seem to reco gn ize th e a rea
Round and round the dusty margin the was p searched
anx i ously and nally ventured to cross the area ( s ix inches
i n diameter ) from which the dust had been removed peeped
once i nto the hol e w hich had fo rmerly been hers and scuttled
away f rantic During he r absence we replaced some dust

arou nd the hol e ; still she d i d not recognize it no r her sp i der
one i nch d istant She cam e w ithin a few inche s o f it on
seve ral occasions but always searched ove r the dusty p er i
h
r
e
and
avoided
the
open
area
w
ith
the
arti
cial
hole
p
y
exactly where he r own had been We nally placed the
sp ide r on the dust i n the region where s he sea rched most
S he returned to the fam i l ia r spot where s he found i t looked
at i t f or a moment but showed no particular agitation
dr o pped her excited man n er walked straight to the hol e
and began d igging i ns ide i t w ithout a moment s delay and
as calmly and busines s like as i f nothing had gone w rong !

A fte r di gging o ut a neat l ittle p ile o f di rt s he came and
.
B EHAVI O R O F P O M P ILI D
WAS P S
47
examined her sp ide r once more pulled and turned it around

legs up and went back to her digging P resently S he re
turned again to the spi der and proceeded to b ite it on the
ventral s ide o f the body between the second and thi rd pai r
o f legs ; we could distinctly s ee her palp i move and we sus
h
s
h
ec
t
e
d
e
that
was
sucking
the
j
uices
When
s
e w ith
p
drew a fter about a m inute s he sto o d licking her chops

repeated the pe r formance and then l e ft promptly w ith such
a strange ight o f abandonment that w e doubted i f she
would eve r return
We in spected the sp ider and sure
enough the body wal l w a s broken at the point mentioned
and a large drop o f clear j uice had exuded
The morning passed and w e saw no mo re o f her
Whether s he became di sgusted with the hard soi l or o ffended
at the di sturbance w e know not B ut it w a s interesting to
see how s he was determ ined to get some good out o f he r
sp ider by at least having a meal be fo re abandoning i t
The tiny parasitic y had been h over i ng over the hole
not over the spider dur i ng thi s enti re per formance and
w a s still poi sed in the ai r over the burrow when we le ft the
scene long a fte r the wasp had given up the enterprise
The spider was taken home At 1 0 o clock that even ing
it responded sl ightly to stimulus but by the next morning
i t was dead
On another mo rn ing as we w ere walking across the eld
past a spot f ro m which all the grass had been trodden w e
suddenly scared up f rom the edge o f the grass a P trop i c us
S he had been engaged i n carrying a spider Ly cosa fr on di
cola Em femal e [ N B anks ] not quite mature but large r
than hersel f A fter a quarter o f an hour s he appeared
,
on
I t is
a n im a l
p ili d a e
q ues t io n
juic es
f eedi n g
rec o rd a s
far
as
wh
for
up o n
we
P o mp i l o i d es tr op i c us do es
e the r
w hi l
t he
k w
no
R o b ert s o n
n ec t a r
o
f P
f o un d
v a ri o us
m a ny
e rs ,
not
l ive
s p ec i e s
he ha s
tr op i cus comi n g to the o
e n t i rely
of
P om
n ot
ers
'
on e
WA S P S TUDI E S A FI ELD
48
searching vaguely in the general region but most o f the

time s ix to twelve feet distant fr om he r sp ider
S he
sea rched by walking a few inches then hopp ing and ying
a foot o r two occas ionally taking a circle on the W i ng j ust
above the grass tops S he seemed to ga in no headway in
her search s o p resently s he went over to a smal l patch o f
cockleburs wh ich w a s S ix feet f rom the sp ider and hopped
and ci rcled ove r the plants w ith an a ir o f getting her bear
i ngs H er behavio r made us wonder i f she had originally
found the sp i der near these plants and had gone back to
retrace her st eps Then s he started out i n the d irect i on
o f the grass plot but mi ssed her p rey so S he went back
to the coc k leb urs again repeated the per formance ; and th is
time went almost directly t o he r sp ider S he po unced upon
i t v iciously as i f s he thought i t necessary that i t be a t
tacke d vigorously and got a good grip grasp i ng i t by its
ventral s i de back o f th e third pai r o f legs
S he dragged i t thus a foot or s o walking backward and
pull i ng l ike a dray ho rse ; then p oised it in some grasses
and went back to hunt a place for a hole S he found a
depress ion in the earth a place where w e had dug w ith a
trowel the day be fore The ground w a s dry and hard wi th
much dust on top S he tried spots here and there i n the
S ide wall s o f thi s hol e scratch ing and b iting fur iously in
e ight di fferent places but s he found the ground too hard
and th e su r face dust fel l i n on her A fte r try i ng thus a
long while S he w ent straight back to her sp ide r tw o feet
then returned to dig as i f
a way and b rought i t nearer
s atis ed that th i s place wa s as goo d as S he could expect to
nd Then S he seemed suddenly to deci de upon he r spot
and a fter j ust a few strokes o f digging for the rst time
s h e swept the di rt back from the space immediately i n front
o f the h ole It was hard digging s o wh il e s he was gone
v
i
i
sp
ide
r
agai
n
we
p
ierced
the
rm
sur
face
crus
r
t
e
o
h
s
t
t
,
B EH AVI O R
OF
P O M P ILID
WAS P S
49
V
A
8
P
E
R
8
0
3
.
8
:
w
e
n
m
c
w
u
w
e
r
e
A
s
a
w
.
0
.
o
5
to help start her nest A fte r th i s she dug rap idly c o m ing
out and v i s iting her spider s ix times wh ile i t lay in th is
position We do not know i f s he exam ined the sp ider f re
quently in o rder to s ee i f it w a s behav ing p roperly or to
gai n an accurate i dea o f the required s i ze o f her hole Next
s he lugged i t one foot nearer to the o p en ing dug out more
earth and again moved the S p i de r neare r to a spot only
n ine i nches from the hol e S he always grasped th e sp ider
by the m iddle o f i ts ventral sur face and carried it vertically
the sp ider s h ind le gs draggin g beh i nd wh ile the front ones
hung l imp ( g 9 ) s he d id not at any time drag it si dewise
by th e legs as other P omp il ids do Furthermore s he always
walked backward when she had the burden and could n o t
s ee her way ve ry well
s o she constantly struggl ed over
obstacles that s he m ight as easily have avo ided i f s he could
have seen where s he was go ing
At one time in moving
her sp ider s he struggled th rough grass and over a tangle
o f weeds for fteen i nches when a p er fectly smooth clear
path lay beside her all the way less than two inches from
her at all po ints
S he seemed now to feel the enthus iasm o f seeing he r
work nearing completion and dug furiously for a few
minutes more ; then s he brou ght the sp ide r very near and
hung i t care ful ly on the tiny weed nearest to the hole only
fou r inches d istant removed a few more mouth ful s and
kicked back the di rt then dragged the arachni d t o the mouth
o f the burrow and attempted to take it in ; but i t was too
large s o she le ft it at the opening o f the burrow and p ro
vigorously kicking the di rt out
c eed ed to deepen the hol e
all over the spi der S he dug on and on i n th i s w ay for an
.
'
An other P
tr op i c us ,
i d er i n p rec i s ely th e
o f i t s v en tra l s u r a c e
sp
up
ab s o l ute ly
s p o ts
w i th
o ut
in
s am e
S he
d i ff e r en t l oc a lity
m a n n er, ho l d i n
al
ay s
p e rp en dic ul a r p l a c e s ,
ev i d e n t
d i f c ulty
w lk
a
an d
ed
it
b ac
e v en
an d
y e a r,
v e rt i c a lly
kw
a rd s ,
b ac i n g
m o v e d he r
b y t he m id dl e
g a l l o p in
u
l ig ht ly
o v erha n
i
n
g g
2
5
not p rot by keep ing her p rey lying at the mouth o f the
hole s o she carr ied it to a spot ve i nches away and pro
c e ed ed w ith he r digging vis iting i t o ften
as i f to s ee that
i t was behav i ng p roperly S he worked by digging up the
loose di rt in the hole and then backing out push ing it up
i n a mound beh ind he r B ut we asked ourselves w hy did
s he dig s o deep and not make the hol e w ide enough to admit
?
the sp ider
T im e dragged b y perhaps S he spent another hour i n
enlarging the hole At last ! f o r the rst time she came
pushing her way head rst up through her mound o f l o ose
d irt O nce mo re s he v i sited the sp ide r and shook it en
i
r
i
n
l
u
by
the
le
ft
leg
w
ent
in
and
pushed
out
the
last
q
gy
bit o f loose d irt emerged head rst again grasped the
spi de r by the le ft c o xa and w ith much labor dragg ed and
tugged i t in Thus w e see that s he made he r exit head
rst only when the nest was rea dy to rece ive the p rovisions
The sp ider tted in the hole so tightly that its legs were all
doubled stra ight back At the last her economy was better
than we had j udged fo r the hol e w a s w ide enough a fte r
all w ith no wasted space and the spider tted i n the hol e
s o snugly that i t coul d not possibly use its legs to kick or
wo rk itsel f free A fter the p rey was in the hol e S he con
t in ued push ing up loose f resh d i rt evidently clearing the
way as s he m o ved it back i n the gal le ry The passage must
have been o f greate r diamete r bel o w the sur face however
i n o rder to allow both her and the d i rt to pass bes ide the
sp ider A fte r ten m inutes s he had thrown up enough fresh
dirt to compl etely close the mouth o f the burrow so we
coul d s ee no more o f he r d o ings S he di d not come out
on the sur face any mo re s ince her p roperty w a s no longer
outside
The quarter hours passed yet S he di d not reappear
Thr i l ling indeed ! Two rational human be ings squatting i n
,
B EHAVI O R O F P O M P I LI D
WAS P S
53
B ut
the burni ng sun watching a motionless spot o f earth
,
how consol ing that the poet who best knew patience has sai d
H e al so serves who only s tands and waits
'
At length we we re p repari ng to dig her up when s ud

d en ly a speck o f the loose di rt i n the middl e o f her mound
quivered and a tiny hole appeared ; s he was hollow ing it
out from underneath and slowly packing it rmly back into
the channel as s he came up until now the door w a s falling
in She emerged gradually slowly packing the loose dirt
back into the h o le w ith all her legs and punching i t down
with the tip o f her ventrally curved abdomen S he lled
it in tight to the top swept the dust back over i t loosely
and departed s o p romptly that we had trouble in inter
c ep t i n g her
When w e dug out the burrow the sp ider kicked so vigo r
The long
o us ly that i t seemed i t would di sl o dge the egg
,
FI G
10
T he b u rro
of
om
p i l o id es tr op i c us
N a tu ra l
si
wh ite egg w a s slightly curved and tted n icely to the side

o f the abdomen where the larva would be in position for
the j uicy tende r part o f its food rst
,
W A S P S T UDI E S A F I EL D
54
The hol e (
g
exact s ize ) one fourth i nch in
diameter went downward at an angle o f 4 0 degrees w ith
the sur face l evel for about t w o inches and terminated in
an ova l h o riz o ntal chamber an inch long and a hal f inch
w i de
From these observati ons w e may be sure that the egg was
depos i ted and the sp ide r entombed at about
a m
At 1 0 o clock that even ing the sp ider w a s active and would
make the characteri stic threatening sp ring back when
touched The l egs could mov e actively but not coordinately
and were un t for progress i o n ; the mand ibl es attempted to
close but could do no more than qu iver when tickled with
a straw A fte r thi s activ ity w e were surprised th e next
mo rning to nd i t dead B y the follow ing day we we re
quite certain that the egg too w a s dead
One m id S eptember morning w e s a w a lot o f d i rt thrown
out o f a hole in the s ide o f a dep ression made by a ho rse s
hoo f As w e came neare r a P trop i c us emerged and ew
away i n alarm but slowly returned found the spot and en
Thus S he ed i n a ffright and tim idly returned three
te red
times when w e tried to come near enough to ob serve her
act i vities
S he j umped i nto the hole wh ich then lacked
o nly a hal f i nch o f being full b it away the soil from around
the edge o f the i rregular hole and dropped it in S til l the
wasp w a s so nervous and susp i c i ous that only w ith the
greatest caution coul d w e get near enough to s ee what s he
was doi ng S he o ften ew out frightened then nervously
ap p ed her w ings walked i n aga in and continued her task
At last w e could creep close enough to s ee her smoo th i ng
down the ll ing i n the hole w ith her abdomen but i n a
strange w ay qu ite new to us The abdomen w a s curled
under the body making the dor s al sur face o f the tip appear
ventral ; th is convex portion pressed against the ground
p roved a very e fc i ent sad i ron smoothing and at the same
time compress ing th e loose soil
s ee
1 0,
B EHAV I O R
O F P O M P I LI D W A S P S
55
Whe n the ll ing w a s packed into the hole up to w ithin

one eighth inch o f the sur face level she kicked i n some
dust from the tiny embankment overhanging the nest not

the yell ow dust that had o riginally come out of the hole
The
sur
face
earth
here
was
grey
and
the
subsoil
yellow
)
(
S he then kicked al l o f the l ittle clods o f grey earth i n a
heap on top went a few inches away and made anothe r neat
p ile o f exactly the same kind then kicked more loose dust
ove r the hole compl etely covering it and also ung i n any
stray lump s large or small that lay near When they were
too large she carried them in her j aws and loosely dropped
them on the site o f the nest At no stage did she attempt
to compress the mass on top w ith he r abdomen ; i t seems
that this treatment i s reserved for interior work Just when
we we re happy to think that the two p iles o f g i ey clods
were placed there to dece ive any enemy that m ight come
near P trop i c us as i f f orgetting he rsel f k icked hal f o f
the second pil e ove r the h o le s o all notions o f actual ra
t io n a l deception being practi sed by th is P ompil id were S hat
-
'
tered
When she cons idered he r wo rk nished she ew to a s pot

fteen feet d istant There she del iberately and restfully
made he r toilet f o r ve m inutes then ew high in the air
and away
We opened the burrow and f ound it as illustrated ( s ee
g
with the bottom o f the chamber three inches belo w
the sur face o f the ground The maximum diameter o f the
chamber w a s one inch The sp ider there in was very active
moving the legs and mandibles violently and giving good
prom ise o f recovering from its abusive treatment The
white egg adhered to the dorso-lateral sur face near the
basal part ; i t neve r hatched Eleven days later the sp ider
L y cosa c aroli nensis Walck [ C R Shoemaker ] was dead
During the same week two P tr op i c us in company with
,
W A S P S T UDI E S
6
5
FI G
1 1
N a tura l
An o the r
si
ze
fo
of
b u rro
A F I ELD
m a d e b y P o mp i l o zd es tr op i c us
two individual s o f a much smalle r species o f P omp i l oi des

were discovered out foraging We watched them for an
hour o r mo re as they walked al ong o n the ground somewhat
nervously exam ining with particular care the base o f all
clumps o f short grass
A month later we chanced to see a P trop i c us foraging
m o re actively S he alighted boldly upon a large spider web
i n the gras s ( one o f the sp reading kind with a den o r lai r
,
'
WAS P S
B EH A VI O R O F P O M P I LI D
57
in o n e corne r ) walked over it with wonder ful lightness and

rap idity and entered the lair S he came out at once with
out the sp ide r however walked w ithout the slightest en
tanglement to the edge o f the web and ew o ff P erhaps
the sp ider spi ed her and escaped but it was surprising that
the P o mp i l oi d es w a s not ensnared in the web as s o many
o f the larger i nsects are especially s ince thi s i s the whole
pur p ose for which the w eb i s constructed We have el se
where remarked that dead Chal y bi on c ae ruleu m and Eu
m en id s are o ften found in spiders webs
Others were seen in other regions and at other times a p
ren tl
a
fo
raging
Some
were
out
in
the
grassy
elds
and
p
y
othe rs in barren clay covered areas They seem to be gen
e ra l ly distributed
whereas B emb ix n ubi lip en n is and othe r
wasps are found only i n speci c places
It seems from what w e have noted that the primary
sting i s insu fcient to paralyze the S pider and that whenever
P trop i c us i s aware o f any activity on the part o f her p rey
s he re stings it
One record shows that a sp ider may recover from the
sting A smal l femal e P tr op i c us i n alarm le ft her sp ider
N
a medium sized specimen o f P ell c n es c o eca tus t
[
B anks ] at our feet on June 1 2 1 9 1 6
Thi s tribute w a s
cheri shed even though i t appeared to be dead
I t was
daily stimulated but i t w a s not until four days later that
the stimulation brought f o rth the slightest movement o f
the legs Every day the sp ider was mildly stimulated w ith
the same sort o f response but i t w a s no t until the fteenth
day that the sp ider sho wed tendencies to revive On that
day June 2 7 i t m o ved the legs more actively and j umped
slightly On July 4 i t was so l ively that when the box was
opened it j umped out I t had also j ust spun a web in the
box The next examination July 1 0 found i t very active
as did also the i nspection on July 2 0 so it was released in the
.
W AS P S TUDI E S AF I EL D
8
5
garden apparently fully revived and capable o f carry ing on

all the duties o f s p id erdo m O f course we must not forget
that the sp ide r rev ived only a fter fteen days and had he re
mained buried w ith the wasp s egg upon h im he would have
been devoured long be fo re the days o f hi s resuscitation
The uid o f the sting must have had some preserving
inuence because without be ing stung it i s doubt ful i f the
sp ide r would have l ived ve weeks w ithout food under
l ike condition s
,
P o mp i l oi d es
m a rgin a tus Say [ 8 A Rohwer ]

.
We followed thi s l ittle black and orange spotted P ompil i d

on the ground for a hal f hour s he appeared to be fo raging
S he kept close to the ground amid the sparsely growing grass
and only once or tw ice cl imbed the g ra ssb la des f or a few
i nche s The tw itching w ings showed that s he might eas ily
be aroused to ange r although the head w a s kept close to
the ground in a very intent manner o ften mov i ng from
s ide to side as i f i n search o f someth i ng A t one time i n
particular i t seemed certai n that s he could n ot be bent upon
any other m iss ion than the ques t o f p rey for when S he
came to the small turret o f a young sp ider s he became v ery
much agitated and l ingered about the place for some time
poss ibly wa iting for the sp i de r to come out and several
times violently ramm ed her head against the turret
Fi
nally seeing the f ru i tlessness o f he r wait here s he continued
3
her wanderings Hancock records nding P m a rgina tus
in the tube o f the castl e bu ilding sp ider L y cosa d om if ex
An hour late r w e watched anothe r o f the same species
going through exactly s im ilar fo raging behavior
We found one on the sunny a fte rnoon o f August 3 1
-
En t
N ew s
1 0 : 29
899
6o
o f P hi dip p us s p [ N B anks ]
She munche d and sucked
at it as i f heartily en j oy i ng her meal
H artman describes the nest building o f P m a rgi n a tus

i n thi s w ay S he had chosen for the home o f her o ffsp ring
the m iddle o f a much used path through the sandy wood s
H ere s he began to di g w ith v im and i n a few moments

had dug a hole an inch o r more i n depth and was bringing
out sand at regular interval s which increased in length w ith
the increase i n depth o f the nest The sand wa s pushed up
in loads with the hind l egs and the end Of the abdomen
The wasp di d not appear w ith a load each time but o ften
ve or s ix loads woul d be allowed to accumulate at the en
trance when the whole p ile woul d be pushed out and scat
t e red away from the entrance more or less care fully

We di scovered another one Octobe r 1 0 j ust in tim e to
witnes s i t do ing a marvellous feat o f strength S he was
walking backward dragging a spide r a Tr oc hos a av ar a
Keys [ N B anks ] much larger than hersel f S he was hold
ing the sp ider i n a vertical position grasp ing it by having
her mandibles inserted in the ventral sur face o f the sp ider
betwee n its l egs I t was remarkabl e that the l ittle wasp
could move such a huge v ictim at all ; but when s he pro
c eed ed to drag i t up a bank eight feet h igh
part o f whi ch
was actually vertical and a small portion sl ightly overhang
ing we could hardly bel ieve our eyes and cannot yet under
stand how i t was possible ( see g 1 2 ) yet the l ittle wasp
seemed not at al l disconcerted about it
Thi s method o f transportation o f prey as illustrated
does not seem constant While Hartman says the wasp
grasped the spi der by one o f the coxae wh ich would prob
ably make the carrying pos ition very nea r to that il lustrated
the P eckhams describe one as going backward carrying
a med ium sized sp ide r

and another dragging a small
.
B ul l U n iv
.
T ex 6 5
.
2
5 54
1 9 05
P O M P I LI D
B EH AVI OR
FI G
12
d ra g g i n
A m a rv e llo u s
he r p rey
up
h i ll
f ea t
of
WA S P S
s t r e n g th :
T w o ti m e s
n a tu ra l
P o m p il o i d es
si
61
marg in at us
62
Thomi sid
the sp ider w a s s o small that S he hel d it in
her mandibles well above the ground and we speak o f her
as dragging i t only because s he walked backward and acted
as though s he were obl iged to exert he rsel f Q
uite o ften
the sp iders taken by this species are too large to be carried
and then i t i s necessary to drag them ; th is h a bit i s s o i n
gra ined that when it would be much more conven ient to go
straight ahead they stick to the ancient custom and seem
unabl e to m ove i n any other way

On another occas ion
they found that s he se ized the sp ider by one leg and dragged
i t o ff and again : We have tw ice seen a m argin a tus p ick

up her sp ide r and y w ith i t backward fo r a long distance
as much as four o r ve feet

When we s ee the constancy o f some wasp s i n the w a y o f
carrying thei r p rey such a s P ri on onyx at r a tu m P th om a e
B emb ix n ub i lip en n is A mmoph il a p i c tip en n is O dy n eri s dor
sal e and others each o f wh ich species transports its pr ey
unvaryingly according to the xed ways o f the specie s and
then w e se e the vari ety o f methods o f P margina tus one
can only wonde r i f natural selection w i ll some day el im inate
the l ess favo rable methods
The sp ide r guring in th e above incident was taken home
When exam ined the next day i t appeared enti rely una ff ected
Its abdomen
b y the sting b ut j umped up and walke d away
had been bitten i nto and the j uices removed s o it was not
at all surp r i s i ng w hen on the th ird day the sp ider was
dead Thus i t would appear that margin a tus sometimes
indulges in animal food although Robertson nds that i t
frequents the owers o f S oli d ag o nemor a lis We have
o ften noted that the p rey o f the P o m p ilo ides exhibit a goo d
deal o f energy and activi ty a fte r they recover from the rst
shock o f the st i ng and soon a fte rwards succumb suddenly
H artman nds that the spide r i s stung to death wh ile the
P eckh ams recor d th at the sp ide r o f t hi s w a sp w hic h they
,
'
B EHAVI O R
OF
P O M P I LI D
WA S P S
63
dug up a fter two days w a s al ive as wa s shown by the

quivering o f the legs ; four days later th is quivering was
They also
s carcely perceptible an d it died two days a fter
mention one instance where the sp ider was stung to death
The one characteristic that i s constant for both H art
man s and P eckham s wasps i s thei r method o f concealment
o f thei r p rey in among the blossoms unde r a lump o f earth
or on a lea f high ab o ve the earth whil e the nest i s being
dug and the hab it o f vis iting i t at short inte rvals while the
wo rk i s in progress
One inte resting reco rd o f the P eckhams w e beg leave
to q uote since it seems as though practical j udgment o r some
othe r intellectual faculty must have been i nvolved
The sp ider was h idden and then began the usual hunting
per fo rmance wh ich soon resulted i n the discovery o f a cav
ity which had a very small opening The wasp crept i n re
mained a m inute and then came out and brought th is spider
to its new hiding place The head went in easily but it
took a great deal o f tugging to get the rest to follow At
last both sp ide r and wasp we re out o f sight and everything
remained quiet for a lon g time When the wasp came
creep ing out s he went on an extended tour She doubtless
selected ano the r halting place fo r when s he returned it
was to try to get the spide r out o f the hol e by pul ling at
one o f the hind legs The task howeve r was not an easy
one ; she exerted al l he r strength s o that they expected to
At
s ee the sp ide r torn to p ieces but still it did not come
last s he seemed to real ize that there w a s more than one way
to accompl ish her end and turned her attention to cutting
away the earth to make the open ing larger A fte r a few
minutes work s he tr i ed again and although the passage
was still much too small fo r conven ience the spi de r was at
length dragged forth
,
Lo c
c it
0
5 .
64
P s a m m o c ha r es
s c el es tus
C ress [ S A Roh w e r ]
.
On October 7 thi s wasp was fo und carrying an immature

sp ider a D o lomedes s p [ C R Shoemaker ]
S he d id not
hold it vertically but drew the sp ider s m iddle leg con
v en i en tly unde r he r body grasped its tarsus in her m a nd i
bles and w hile s he walked forwa rd dragged its b od y along
bump ity bump over sticks and stones A fter travell ing th is
way f or a short time s he h id the sp ide r under a clod and
ew about over the sandy bank fo r some l ittle time where
she p robably found a suitable spot or hole ; then S he returned
to her sp ider and again in the same way attempted to pull
i t up hill Thi s time howeve r s he failed to get a rm
foothold and both rolled down to the l edge below H ere
s he carri ed i t to a small dep ression and le ft i t
When s he
failed to return a fte r an hou r we took the spider which
B y the next day it had regained its
w a s then motionl ess
Vigo r to such an extent that it coul d sl ightly move the body
si dewi se Three days a fter its capture w hi le it coul d not
move its body th e legs responded to stimulus The next
day it w a s so active that i t could hop for some d istance w ith
surp rising agility but on October 1 3 s ix days a f ter its
capture i t w a s dead
One day i n searching fo r insects w e dug a l ittle cave
seve ral inche s deep into a bank near its base Two days
late r October 5 this black wasp w a s at work digging a
hol e at th e extrem e end o f th is l ittle dugout The hole
must have been deep o r large fo r already there was a
goodly p ile o f fresh earth thrown out about the or i ce It
was rathe r dark back i n thi s l ittle cave but with the ai d
o f l ight reected upon the wasp f rom a m irror w e we re
able to follow its maneuvers The wasp went int o the hole
head rst remained inside a good many m inutes and when
.
T hi s
a sp
i s th e
s am e a s
omp il u s
s c e l es tus
( d e R o hw er )
B EH AVI O R O F P O M P ILI D
65
WAS P S
nally she appeared s he brought w ith her under her body

an astonishingly large load o f dirt One coul d easily spell
o ut her method : during her long stay in the hole s he loo s
ened the so il and kicked it under her body then on coming
out o f the hol e backwards s he had only to spread her fore
legs in such a way as to make a fence or rake and by pass
ing out S he woul d automatically drag out the loose soil
When one sees the slope o f the burrow i n g 1 3 one
,
FI G
iz
e
T he b urro
13
and
rey
of
s c c l es tns
N a tura l
real izes her tremend o us task in pushing he r load up hill

back wards S ometimes the l o ad w a s s o large that the wasp
had to force he r w a y w ith much e ff ort out through her
doorway Occas ionally the loose earth at the top o f the
-
66
gallery w a s kicked out but no attempt w a s made to clear

away the d b ris f rom he r own doo r
We looked for a sp ider which we suspected she must have
near by but found none Once when we intruded too close
s he ed to the top o f a bank some fteen feet away
but
soon returned to he r hole A fter a few m inutes more o f
digging s he ew again to the same place on the top o f the
bank and p icked up the prey S he had de ftly concealed among
the vegetation S he at once grabbed i t by its ventral cen
te r whe re the legs j oin the body and walked backward
toward her hole w ith the sp ide r i n a vert i cal pos iti on S he
reached the ledge j ust above the hole dropped he r sp ider
and entered a fal se hole but soon d i scovered he r error took
up her burden and p roceeded di rect to he r own burrow
leav i ng the sp ide r twi ce wh ile s he went on t o exam i ne the
n est ; but on these last two v is its s he did not stop to get
a rm hol d upon i t a s be fore but grasped it i n any way
convenient by the head o r legs to drag i t to the hol e At
the very brink S he lai d her V ictim down fo r a moment
w i th its head toward the hole wh il e s he went ins ide for the
nal inspection S he came up and slowly and del iberately
turned the sp ider around graspe d i t by the anus and
dragged it in As i t w a s pulled thus into the hole all the
legs turned forward and p ressed rmly against the body o f
the sp ider ; then w e coul d s ee the logic o f her choos in g that
pos ition B ut what impressed u s most w a s the del i b e rate
and care ful manne r which the wasp di splayed in turning
the a rachni d around so that the rounded and a p p en da g el es s
part could be grasped I t was no accident but a del ib
e ra t e act
She remained with in for one hour When we opened the
nest w e found the sp ider resting still with its head toward
the exi t ( g 1 3 exact s i ze ) and on its ri ght s ide w a s the
wasp s egg The wasp s last hou r had evidently been spent
,
68
head probably ve time s a s heavy as hersel f an d was walk

ing backwards dragging i t ove r sticks and stones green
and d ried grass up l o gs and down
A fter four minutes the wa sp stopped dragging the sp ider
and for ve m inutes she rested and cleaned her antennae
body and legs Twice during thi s interval s he le ft h er sp i der
and went under a lea f near by but each time she returned
to keep watch aga in i n f ront o f its prostrate form wh il e
B ut w e were m istaken i n thinking
s he made her toi let
that her burrow w a s under thi s l ea f fo r at 1 :2 9 s he grabbed
her p roperty and continued her j ourney i n the same manne r
a s be fore for s i x m inutes
Tw ice s he l e ft the sp ider
walked about in ci rcl es w ith the same nervous j erky gait
but in a moment she se ized i t and hurried on agai n mak
ing good time ove r a b it o f smooth road which chanced
to be in he r w a y S he ce rtainly seemed not to trouble her
sel f to nd a smooth w a y One o f the spide r s legs w a s
seen to quiver several times
At
the wasp enj oyed a per fect rest near her prey
for a hal f m inute The sp ider s second right leg still q uiv
e red occas ionally ; but the wasp was not perturbed by it but
calmly rubbed he rsel f d o wn
Sh e cleaned her abdomen
w ith he r hind legs and her antennae and head w ith the fore
legs Fo r nine m inutes s he lo itered about mak i ng he r toilet
most o f the tim e and occasionally mov ing her position for
a few inches sometime s behind o r to the right o r l e ft o f
her sp ide r but always near enough to keep a watch ful eye
upon it
The sp ide r s r i ght leg gave a more marked tw itch The
wasp darted at once to her position at the right o f the
spide r s head At 1 :4 4 she le ft i t and went under a dri ed
lea f as i f reconno i teri ng A fter a few seconds s he came
out and dragged h er p rey under the lea f w ith her the
legs o f the sp i der p rotrudin g We f ollowed eagerly but
,
B EH AVIO R O F P O M P ILI D
WAS P S
69
cautiousl y thinking that here s he woul d bury it B ut no

s he could b e seen
through a rent in the lea f quietly at
re st be fore her sp ider The wasp occas ionally walked about
slowly under the lea f ; apparently s he only wanted a mo re
sheltered spot wherein to rest and loiter Up to this point
s he had travelled about sixty feet
Two o clock came and there was no change ; the sun beat
down hotly and our patience w a s strained With the for
ceps w e gently pulled the spider out by its leg fo r a hal f
inch
Th is instantly aroused the wasp to action ; s he
snatched the spide r by i ts head and d ragged it under the
lea f clear to the extreme opposite margin paused a mo
ment as i f undecided and then impetuously se ized her
p rey and resumed her j ourney in earnest There was much
di fculty in following he r owing to obstructions vegetation
and poi son ivy S he showed by her actions that she now
felt anxious and skeptical i n our presence ; from
to
2 :1 2 she remained at rest but this time holding the sp ider
in he r j aws all the time Finally s he regained enough con
d en c e to lay i t down wh ile she made he r toilet for three
minutes We should b e glad to kno w i f s he purpo sely laid
down her p rey so o ften in order to clean hersel f or i f she
only rubbed hersel f fo r a pastime when s he paused to rest
We have also wondered i f all th is was for the purpose o f
personal daintine ss o r i f a part o f thi s practice served as
a massage o r an athleti c rub down o f the ti red muscles a fter
the strain o f labor
At 2 :1 5 the wasp made a very grace ful attempt to cl imb
a log which lay over her path H ow the insect became aware
o f its presence w ithout facing i t to see it or bump ing into
i t w e cannot imagine but she seemed somehow to sen s e the
l ocation preci sely S he backed up nea r to it dragging her
sp ider until she came within reach o f the lo g when s he
l i fted her hind pai r o f legs h igh up in the air until they
,
A F I ELD
WAS P STUDI E S
7o
'
came in contact w ith the rough bark ; then s he c on t in ued

backing up unti l s oo n s he had all s ix o f her legs i n action
Just a s s he reached the top w ith her heavy burden howeve r
S he grasped a f a llen l ea f which lay loosely upo n the log
and l ea f wasp and sp id en ro lled to the ground F i nding
her l ost p rey a fter a few seconds search the wasp resumed
he r travels in another di rection thi s time up a steep hill
s ide
S he resorted to no diagonal i ncl ines but pluckily
struggled stra ight up the steep slop e paus ing only once
fo r a few minute s re st when hal f way up the hill P res
ently she sl ip p ed and fell and rolled part way down the h ill
This seemed to discourage her a little and she le ft her p rey
and strolled a dozen inches away th en back to it thought
fully and settled do w n to spend the next ten m i nutes i n
resting toilet making and S hort j erky walks
At last s he resumed her wanderings ca rrying the sp ider
still further up th e h ill and reached an area blackened by
a recent re where there was no green vegetation and the
ground w a s l iterally full o f the holes and tunnels o f ro dents
B ut nothing he re seemed to appeal to her and aga in she
took up her burden and went down the hill beyond for ten
f e et and then up the next h illock always walking back
wa rds and hold ing the sp ide r by i ts head About thi s time
she dropped he r sp i de r two or three times and made short
s ide trips looking ove r the v icin ity return ing to i t a s though
to continue the tri p S uddenly and w ithout warning s he
disappeared w ith he r p rey into a ro dent hol e We watched
and waited long and wasted much e ffo rt i n digging up the
tunnels and a considerable portion o f the surrounding coun
try but could d iscover her no more
Over the very roughest country thi s wasp had dragged
he r burden fo r a hundred yards S ince she s eem ed to have
one w onders w hy s he
a n especial l iking fo r rodent holes
d id not use f o r he r nesting place the rst tunnel where in
.
B EH AVIO R O F P O M P ILI D
WA S P S
1
7
she found the spider and thus save hersel f the labor o f
dragging such a load for three hundred feet over rough
roads or at least choose one o f the many which S he passed
by in the burnt area
A fter following this wasp mothe r through the details o f
her prolon ged trials and di fculties we respect her fo r her
faith fulnes s and pluck in continuing her heavy task to the
end And yet we have a lurking feel ing ( wh ich we regret
to express ) that this type of faith fulness may have been
an exhibit more o f brawn than o f brain Howeve r we
must not j udge the species by thi s one female which may
have been very ind ivi dual istic in her behavior ; perhaps her
s isters would have shown as ne instinct and economy as
i s found in any o f the other species
,
P ri o c n em is p o m p i l us
.
Ver i ly i f boil s were sent to try the patience o f Job then

P ri o cn emis p omp ilus must have been created to try the
patience o f natural ists On e sees an an imated l ittle black
wasp darting nervously here and there in a hot sandy eld
and one follows eagerly c o n den t that some interesting
activity will ensue at any moment S he runs in this dirce
tion and that absolutely w ithout any system or general
d irection peep ing into every crevice or beh ind eve ry ob
o ften hopping and twitching the w ings nervously
s ta c l e
Thus s he lures us on and on almost interminably until our
eyes feel strained w ith watching her in the glaring sun
Sometimes she plunges into a crevice o r a mole s tunnel
and w e wai t until she come s out empty handed or s he may
walk about the tunnel and emerge far beyond and leave us
watching for ever s o long at the place wh ere s he entered
,
T hi s s p ec i e s
( de R o hw e r )
7
i s l i s t ed i n
D a lla T o rr e s
C a t a l o gue
as
S a li us
p omp il u s
2
7
be fore w e discove r that w e have been dece i ved scramble

to o ur feet and go scurrying a fter her again O r perhaps
and
s
its
s he dar s under a grass blade and there s h e s its
t
and sits occasionally scratching a bit at the ground o r peep
ing out as i f to make sure that her secrets were not being
sp ied upon and eventually darts out w ithout doing a thing
and resumes h e r crazy j ourney P robably these are f o ra g
i n g trips but in the summer months we have never seen
the i r activity lead to anything more than thi s vain search
It may be because o f the scarcity o f sp iders o f adequate
siz e at that time o f year o r due to the sexual immat u rity o f
the wasp s Late r i n the season S eptember and October
the ir work seems to be mo re purpose ful and thei r e fforts
lead eventually to some results
On the a fternoon o f October 3 1 9 1 6 we met w ith one
such i nstance A P ri o c n em is p omp ilus was rushing hither
and thither ove r a small area o f broken gr o und We had
so o ften seen thi s speci es wande ring about on the bare
ground inspecting crevices and had been s o co nstantly d is
appo inted in seeing them do nothing that we paid no atten
tion to th i s one Upon returning a n hour later howeve r
and nding apparently the same wasp runn ing about in the
same small a rea we suspected that there must be sp iders
in the many depress ion s i n the earth so we sat down and
wa ited
The retina o f one s mind a s well as the retina o f the eye
grows dull when weary o f gazing at one thing so our
thoughts were far away when pe rhaps a hal f hour late r
suddenly there was a great commotio n hurry tumble

topsy turvy ! The wasp had her spi der and was stinging i t
The sh o ck was so great and the w o rk was d o ne so quickly
that we could not s e e j ust how i t w a s done The wasp le ft
he r V ictim ran around l ike w ildre for several seconds
returned stung it again and dragged it s o me di st a nce away
,
WAS P S
73
le ft it and in the same w ild manne r looked f o r crev ices

went fro m one to another and eventually carried the sp ider
Not sati s ed w ith thi s the wasp dragged
to a shallow one
her burden further and in an uncertain way wandered
ab o ut w ith it P resently she l o st her prey but c o ntinued
r o aming ab o ut possibly looking for it
B e ing unable to wait longer we took the wasp to make
sure o f the spec ies Whil e the story here i s incomplete
these observations S how that these wasps are p ersi stent in
remaining in a spot when they are susp icious that thei r prey
l ies hidden thereabouts and that they get thei r prey rst and
then hunt for a suitable crev ice wherein to deposit it
Even later in the season than thi s October 2 6 the active
P p omp ilus were more abundant than at an earl ier date
Wh ile these wasp s had frequently been seen be fore in search
o f game on thi s day f o ur were di scovered with sp iders
on a clay bank
One in particular was followed and
watched
S he had a medium sized S p ider i n her mouth
and w a s ne rvously and quickly moving w ith i t We say
moving because her gait ha s no name ; she was walking and
ying w ithout l i fting he r legs from the ground and doing
both equally well eithe r pushing her burden forward o r
going backward and dragging it w ith her S he would o ften
leave her prey to explore the region f or suitable holes At
last one was found o n a pe rpend icular bank e ighteen inches
f rom where th e spider lay Thi s bank was alm o st straight
up and down and w e wondered how it would b e p ossible
for the wasp to carry the prey up to the hol e It seemed
fo r a time as i f i t were not possible fo r in the twenty or
more attempts that s he made walking backward and cl imb
ing up the bank invariably both fell down ; e ithe r S he
tumbled down hold ing to her prey o r s he dropped i t and
ew d o wn a fter it She made a few exam inati ons o f other
n earby h o les
as i f cons ide ring the advi sabi lity o f a change
,
W A S P S TUDIE S AFI ELD
74
o f plans but for some reason she always retii rn ed to the

one o f her rst cho ice
Eventually s he mastered the art needed and her persever
ence was rewarded with success ; at last S he got the sp ider up
to the hole H oweve r the l edge i n front o f the hole was
s o narrow that i t would tumble o ff j ust as fast as s he got
i t there I t seemed that s he had an avers ion t o dragging
the spi der i nto the hole w ithout rst going in he rsel f and
thi s seemed impo ssible S he late r learned to place o r bette r

t o s a y succeeded in placing the sp ide r j ust over the mouth
o f the burrow and there i t hung whil e s he very dextrously
wedged her w a y into the hole to s ee i f all w a s right be fore
entering w ith the p rey B ut here new disaster came ; s he
could let the sp ider lie on the outs ide ove r the hole whi le
s he squeezed hersel f in
but when s he came out he r head
butted aga i nst the sp ider and down i t went again int o the
valley below H e r patience and strength seemed i nexhaust
ibl e w e had long ere th i s lost all estimate o f the number o f
times s he had skil fully carried he r sp ide r up this p recip itous
bank Three times in succes sion th i s last m ishap occurred
when nally ( and thi s may have been accidental but we
shall always hol d i t as a case o f p rot i ng by ex p e rience )
s he took up he r spider
carried i t some distance to a po int
where the bank sloped gradually and cl imbed up walking
backward and pull ing the sp ide r with he r ; then she dragged
it back horizontally and let it rest on a l edge three inches
di rectly above the hole Next s he went i nto th e n est ex
H ere
a m i n e d i t once more and came out and got he r booty
we thought the insect showe d wonder ful intell igence ; it
woul d be eas ie r to carry h er sp ide r down three i nche s to
the hole than up e ighteen i nches and still it would be w ith in
easy V iew wh ile s he w ent for her nal inspection o f the
hol e wh ich s he deemed s o necessary Th is arrangement
w e thought woul d obviate that exam inati on and the s ub s e
quent sl iding down o f the sp ide r into th e chasm below
,
WA S P STU DI E S
6
7
A F I ELD
hour s wea ry waiting fo r the wasp to eme rge w e st i rred

he r out w ith a p robe and dug away the earth to explore
the nest I t went in hori zon tally about s ix inches ( it had
p robably been a beetle s hole ) then the channel o r crevice
was lost The prey wa s not found P robably her last hour
was spent in ll ing and packing the hole with in up to thi s
i
n
m
P
h
a
fter
the
manne
r
o
f
other
omp
il
ids
mentioned
o
u
t
c
p
Thi s species o f wasp uses a variety o f spiders We have
at va rious times and places found it carrying o ff the follow
ing species : P ar d os a m i n i m a Keys [ N B ank s ] s p o f Ly
i mmature [ N B anks ] L y cosa helluo Walc k
c o s i d ae
B
o
P
young
anks
P
r
s
a
n
i
r
l
i
Em
c
n
a
i
N
a
d
a
s
a
]
g p p
) [
(
Tl ]
UH
FI G
p
r ey
14
P ri o c n em is
T w ic e
d en s is B lk
B anks ]
n a t u ra l
[N
p omp il u s
s ize
tr a v elli n
g b kw
ac
a rd s a n d
d ra
gg i g
n
he r
B anks ]
and Cl ub i on a
b b o tt i Koc h [ N
I t s oon becomes apparent that i n many o f the a ff a i rs o f

wasp l i fe P p omp ilus i s not such a stickle r fo r convention
al i ty a s are many o f he r s i ster species S he takes such prey
as she can get and d oes not rej ect good spi ders because
they are not o f the spec ies wh ich H ab it speci es as p rope r
fo r her ; s he uses such holes as she can nd i n the moment
o f need and mod i es them to suit he r needs ; i f she cannot
ascend a precip itous slope d i rectly s he w ill in time contrive
some indi rect way to scal e it In fact l ike most individual s
who go outs ide the pal e o f convention while s he may at
,
WAS P S
77
times seem bizarre yet her action contains an increased

portion o f re fresh ing multi formity and resource fulness
And s o also in he r w a y o f carrying her prey S he has not
one xed way but a variety o f ways She seems in fact
to be enti re ly without any xed habit in thi s point except
ing that s he grasp s it in he r j aws T hi s i s really the only
way possibl e for her s ince she i s s o small that she could
not poss ibly straddle sp iders o f the size s he catches H er
most common mode o f locomotion is to walk backward
pull ing he r p rey ( g 1 4 ) but S he also sometimes walks side
w i se dragging her burden a t right angles to he r ow n body
H er method o f hal f running and hal f y in g with the feet on
the ground whil e s he e ithe r pushes o r pulls her sp ider has
al ready been described I n m aking ready to drag he r prey
she se ize s it by any o f its members that may o ffer them
selves as a conven i ent handle at the moment Each place
o f insertion o f her j aws in the sp ider s anatomy gives i t a
di ff e rent aspect Fo r inst ance when s he grasps the sp ider
by the base o f one m iddle leg thi s stand s the body on its
s ide ; when s he grasps i t by the m iddle o f the ventral s ide
o f the tho rax the ponderous abdomen bob s up and d o wn
as i t passes over the rough ground in a way that must be
annoying to the l ittl e worke r Anothe r o f her favorite
methods i s to sei ze it by the palpi T he lo os e sand o ften
gives way unde r he r feet and both captor and prey roll
headlong
Another species o f P ri oc n em is ( P a vi c orn is ) according
8
to Needham and Loyd transport their prey by ying above
the sur face o f the wate r and towing the loa d too heavy to
be carried Out onto the sur face o f the water the wasp
drags the h u
ge l imp spide r and m o unting into the air with
her engines go ing and her wings s tea d ily b uz z in g s he sails
across the water trail ing the sp ider and l eaving a wake
,
Li f e
of
I n la n d
W a te r s
330
916
s
7
that i s a m iniature o f that o f a passing steame r S he sa i ls

a di rect and unerring course to the V icin ity o f he r burrow
in the bank and brings her cargo a s hore at some nearby
landing
.
Ara c hn op ho c ton us
ferrugi ne us S ay [ S A Rohwe r ]
1 0
Thi s large red wasp was observed on July 7 1 9 08,

walking up the perpendi cular stone wall o f the abandoned
i ce house at M eramec H ighlands near S t Loui s S he was

walking backwards dragging a very large L y cosa sp ider
much heav ie r than he rsel f straigh t up the wall S he hel d
the sp ider s head between her mand ibles and w a s mak i ng
ve ry rap id p rogress Thinking that s he was cl imb ing to
some height in o rde r t o y down to he r nest ( i n thi s con
r
e
c
t
u
e
we
were
p
robably
m
istaken
captured
her
and
e
w
j
)
he r prey be fore s he should get b eyond reach The sp ide r
had a long gash surrounded by a large swell ing on the
do rsal s i de o f its abdomen but w e had no way o f know ing
w hethe r this w a s a wound in icted by the wasp s sting or
a cut from he r mandibles or only a laceration f rom the
stones over wh ich i t had been f orcibly dragged
Whil e looking for mu d dau b ers nests about the stone
foundation o f another abandoned building i n this v icin ity
we frightened anothe r sp ecimen o f thi s formidabl e looking
species out o f a crevice
The cracks between the l ime
stones o f th e foundation were lled w ith well weathered
mortar S he ew to the ground nea r by and crouched low
un der some grass ; when we attempted to e xam i ne her she
,
M r Roh
.
div i s i o n s
of
T hi s
1 0
er
th e
a sp
r i te s
old
is
co
t ha t th i s
mp a r
en e ric
o mp il u s
a tiv e ly r a r e
g e n us P
m en ti o n e d he r e we have
i n ve y e ars
,
seen
n am e
is
one
of
As hm ea d
o n ly
i n thi s
ve
re g i o n
; b e s id es the f ew
f o ur f em al es
an d
on e
m a l e,
WAS P S
B EH AVI O R O F P O M P ILI D
79
ew further but s he was recognize d immediately by the

bright red o f her abdomen and i n the bright sunl ight her
bl uish tinted w ings H e r brill iant coloration and he r nerv
o us deant manner ce rtainly render her a form idable look
In cabinet specimen s o f thi s species
ing creatur e ( g
the red o f thei r bodies looks far more dul l and the w ings
,
FI G
Exact
1
si
Th e P o m p i li d
asp
Ara c hn op h o c to n us f erru g i n eu s S ay
which are a consp icuous royal blue when V ibrating in th e

sunshine are but a sm o ky colo r with a slight blacki sh blue
tinge at the base
A fter having frightened her away we p roceeded to ex
am ine the loose mortar fo r her nest o r p rey We were
attracted to a p ile o f loose d ry plaster dust on the damp
earth ; thi s po inted up to the thi rd tier o f rock p robably
e ighteen inches from the ground where a large L y cosa
spider lay i n the crevice about an inch from the opening
I t w a s removed and found to be f resh l imp and motionl ess ;
i t was probably dead so we replaced i t and waited A fte r
fteen minutes th e wasp ew into the v icinity again but
bec ame s hy upon seeing us there A fter another ten mi n
utes she ventured near again and al ighted on the stone wal l
at a height o f S ix feet and east o f the nest ; then walked
down and westward entering and examin ing several hole s
and remaining in some o f them for a few seconds S he
eventually found the crevice containing the spider en
tered and examined it so i n tently th a t we w onde red i f she
-
8o
recogn ized that i t had been tampered w ith Then s he turned

he r abdomen toward the spider and held it thus hugging
i t closely fo r some seconds ; we suspected that th e wasp
was stinging it again but she was so deep in the crevice
that we could not s ee d istinctly S he then went deeper into
the cav ity between the stone s and ung out another hal f
dozen kicks o f dust Un f o rtuna tely at thi s point s he sp ied
us lying at on the ground to scrutinize her so s he darted
away to another part o f the wall and entered another n iche
and remained there so long that we thought she would
carry he r sp ider there She suddenly emerged f rom thi s
retreat and escaped We waited an hour for her return
but s he seemed s hy so we wandered away returning every
three or s ix m inutes to look for he r
While thus lo itering about waiting we took a trowel and
at a po in t i n the wall about th irty feet away i dly removed
from the foundation the hard black cement wh ich c o vered
the so ft mortar beneath thinking thereby to make an a t
tractive spot fo r othe r wasp s A fte r a time w e l ost p a
t i en c e howeve r and decided to work elsewhere
We knew
the speci es o f the wasp the place where her sp ider lay w a s
marked and w e knew where she had dug her cavity in the
mo rtar s o i t was decided to l eave he r in peace and return
later to get the spi der w i th the egg on it
Upon returning in a hal f hour imagine our surprise upon
nding the sp ide r gone I t seemed scarcely p robabl e that
another had come along and purloined i t because f rom
her p revious acti o ns it appeared that she was go ing to
change her nesting place Now it was a task to nd the
sp ider and for anothe r hal f hour we dug into the mo rt ar
i n variou s susp icious looking places i n search o f i t S ud

d enly a pass ing gl eam o f color caught our eye ; the wa sp
ew past igno ring her old location and al ighting on the
h
al
h
irty
feet
beyond
We
watched
her
care
fully
as
e
s
t
l
w
.
B EH AVI O R
OF
P O M P ILI D
81
WAS P S
tried to nd her new apartment S he accompl i shed thi s w ith

di fculty ; twice s he had t o y away and return on the w ing
Eventually S he located
t o the same spot to get her bearings
her hole and entered the very one we had only a sho rt
time b e fore begun i n order to attract some wasp ! We were
almost too astonished to believe the story ourselves and yet
the evidence seem s su fcient to warrant the surmise that it
was the same wasp We knew that s he had become d i s sa t is
ed w ith the place that harbored intruders and that no others
o f thi s species were seen al l that day Her actions ind icated
that she was looking fo r a new abode and we are sure that
this sec o nd hole could not have been the re for more than a
hal f hou r the exact time since the rst nest had been de
s e rt ed
because i t had been access ible only since we pulled
away the hard outer cement
She went i n and remained about fteen m inutes when
s he was captured in a V ial placed o ver her hole
We worked
fo r a long time trying to dig out he r spider and egg ; but
the p robe showed that the crevi ce between the rocks was
eight inche s deep s o w e we re o bl iged to give up the quest
w ithout getting he r prey o r learning anything more about
he r nest o r egg
Thi s al s o solves the p roblem o f seven years p reviously
when an A ferrugi n e us ascend ed the o ld st o ne wall o f the
ice h o use walking backward and dragging he r L y cosa N o
d o ubt that wasp al so was di recting he r cours e to a ch o sen
c revice i n the old stone wall
In o ther species o f P o mp il ids as previously noted when
the wasp i s digging her burrow she o ften leaves her work
to exam ine he r prey and f requently moves it closer to the
hol e The notes that f o llow will S h ow that A ferru gi n eus
does not drag her p rey along until she c o mes to a suitable
spot but in tran s porting her prey s he ha s a very de nite
n o tion o f where s he i s going
.
82
On August 1 9 at
p m one was d iscovered drag
ging a large Lycosi d sp ider by one mandible walking back
wards with a j erky motion travelling w ithout much d if
culty over the long fallen blue
grass in the front ya rd She
carried the sp ide r right side up and brought it about fty
feet a fter being d iscovered Tw ice in that time s he le ft i t
and took a quick ci rcl ing ight as i f exploring over to the
ol d log smoke house w ith plaste r ch inked crev ices ; then she
returned without di f culty to her sp ider and co ntinued her
course ve ry di rectly as i f s he knew p recisely what s he was
about When nally s he got it to the bal d area a few feet
in front o f the smoke house s he paused le ft i t again and
ew to the wall al ighted on it and took a qui ck surveying
gallop over the two lowe r logs and the i r interstices ; then
went right back and got the sp ider took it hastily over to
the wall and hi d it deep i n a crevice in the foundation S he
then took on a very ne rvous attitude running and ying
all over the two walls exam ining nooks and crannies every
where wi th w ings and antennae nervously V ibrating ;
na lly s he sat down f o r ve minutes rest and medita
tion then resumed the chase and search We cannot b e

l ieve that the wasp travelled from the sp i der to th e smoke
house s o o ften to see i f the smoke house was still extant
but we th i nk pe rhaps that w a s he r way o f ascerta ining
whether s he w a s travelling in the right d i rection ; she knew
where s he wanted to go and retraced her steps o ften to s ee
that she wa s go i ng there
P resently the w a sp di sappeared in one o f the crann ies
M eanwh ile the sp ide r was exam ined I t was l imp but re
s p o n d ed to stimulation by sl ight movements o f the legs
although it was so completely paralyzed as to be almost
dead
A fter a hal f hour i n hiding the wasp again a p p p ea red
and made he r w ay to the spider and rested i n the crevice
.
'
84
Tr a chelos
tr an q u illa H entz which
much heavier than

he rsel f The sp ider s legs had been b i tten o ff at the j unction
o f the c o x a c and t rochanters
She dragged it w ith its
,
w as
FI G
16
T he t
t he ho l e s m a d e
wi
( is
n -c e l l e
b y the
n e s ts
of
gi g w
a sp
em e r
a r c hi te c ta ,
P s eud ag e n ia
x a ct siz e
s ho
wi
ng
smooth rounded back downwa rd to prevent friction and

grasped i t w ith j aws near the t i p o f the ventral sur face o f
the abd omen
3
Ashmead in citing Walsh and Riley says that the thim
ble shape d cells occur under bark logs o r rocks and are
paras itized by P ter o ma l us s p and Os p ry noc ho tus jun
ce n s C r
,
P s eudag en i a p ul c hrip en n i s
.
Th i s medium S ized black wasp was found walk i ng along

a path at C reve Coeur Lake on Octobe r 7 ca rrying a sp ider
The wasp
P hi di p p us a n dax H entz [ C R Shoemaker ]
w a s holding the sp ide r by one o f its chel icerae and walking
s idew i se the comb inat i on o f sp ider and wasp form ing a
right angle ( s ee g
It moved in a very strange man
ner walking sidewise Thi s wa s the rst wasp we had
ever seen travelling thus N ight was app roaching and tra in
time w a s draw ing near so we were obl iged to abandon
-
1 3
P sy c he 7
66
89 6
B EH AVI O R
OF
W/
nI
FI G
b urro
17
85
WAS P S
P O M P ILI D
P s eud a g en ia p u l c hr ip en n i s
T w o t i m e s n a t ura l s i e
e s c o rti n g
ppu s
P hili
a n dax
t o her
furthe r observati ons on thei r free behav ior and take both
The sp ider showed signs o f l i fe fo r three days
Long a fterward anothe r wasp was found actively en
gaged i n running about on a fallen log Suddenly i t leaped
to the ground and chased a medium sized spide r The arach
ni d made one leap and escaped The wasp became frantic ;
the s ight o f the sp ider and the fact that she had been f o oled
so excited her that she acted as though mad walking ying
running about the ground in indescribable di rections until
A fte r a few mo
s he tumbled headlong into a mole s hol e
ments she emerged from the burrow carrying a sp ider by its
anus S he walked fo rwards backwards or sidewi se ove r the
ground or low vegetation in a manner indicating intense
excitement and eluded us many times be fore we nally
captured her All this occurred not in the ho t sunshine
where most P ompi lids choose to work but on a path in the
cool shady forest
.
86
P s eu dag en i a m ellip es
S ay [ 5 A Rohwer ]
.
We have reco rded and illustrated the fact that P m el lip es

emerges f rom cells made i n the walls o f the mud nest o f
S cel iphro n
The gure here repr o duced ( g 1 8 ) shows
the nest o f the mud dauber w ith two h illy p rotrus ions which
were cut down to show the cells that conta i ned the pupal
.
FI G
18
T w o p ro tub e ra n c e s o n the
o rth a d ul t s o f P s e uda g en ia m ell ip es
.
of
n es t
N a tura l
w hi
S c e lip hro n
si
ze
ch
gv
a
cases o f P m el lip es Elsewhere on the nest mandibl e marks

were i n evidence so we suspect that m ellip es dug out a small
cavity and then carried mud b itten from other parts o f the
nest and built up and around it until the cell w a s completely
enclosed and hidden f rom v iew
Th is we nd i s not the only method o f n id i c a t io n o f
this species At Mosel le M i ssouri on June 30 1 9 1 6 while
b reaking away some loose bark from a fallen tree we found
a l ittle three celled nest ve ry beauti fully constructed wh ich
w e care ful ly guarded s ince we expected some species o f
A g e n i a to emerge there f rom
We were surprised when on
July 8 and 1 1 two adults o f P m ellip es eme rged The nest
did no t adhere to the wood but lay loos ely under the bark
T he accompany i ng gure ( g 1 9 ) shows th i s nest and also
the builde r ( exact s ize )
.
14
o u rn
An im a l
B ehav io r
: 2 7 -63
91 6
W AS P STUDIE S AFI ELD
88
FI G
th e
20
T he m ud
M u ti l l i d
a sp
c el l s
of
P s eud ag en ia
S p ha er op ht ha l ma
f ro m w hi c h
em er
s c a ev a
ed
a dul t s
N a tu ra l
of
Si e
says that A g en ia m ellip es makes tli im b le shaped cells under

bark log s and stones and that they are paras itized b y
P tero m al us s p and Os p ry n o c ho tus j u n ceus Cr
The cell s
here illustrated can hardly be called th imble shaped
A collection o f P s eu dag en i a s p mud cell s was f o und
unde r the loose bark o f a tree ( g 2 0A ) and another
-
1 5
Lo c
c it
66
B EH AVI O R OF P O M P ILI D
WAS P S
89
group o f identical cells was found under a plank wh ich

covered a depression in the ground B oth lots however
.
FI G
20
S l ig ht ly
A m ud
e n la r g e d
n es t o f
P s euda g en ia m el lip es
w i thi
an
o ak - a p
p le
gave us not P s eudag en i a wasp s but both sexes o f the para

sitie cow kille r S p ha erop hthalm a s ca eva B lake [ S A
Rohwer ]
,
CH A P TE R I I I
SOME
H yp o cra b r o
F L!
s ti r
AT CH I N G WA S P S
i
c
o
l
u
P
ack
s
p
[ 8 A Rohwer ]
An old railroad tie lay hal f buried i n a heap o f dirt

Under the protrud ing rotten end w a s a m iniature h ill o f
sawdust or rather particles o f wo od b itten out and above
th i s a clean cut hol e pierced the log We found a fter w e
began digging into i t that only a small portion o f th i s w a s
rotten and unde r the sur face the wood w a s very hard The
long tunnel s l eading in f rom the sur face aperture were
made in the so fte r decayed part o f the w oo d Thi s tunnel
led to a seri es o f three cell s wh ich were ve mm i n diam
eter and were partitioned to make neat cells fteen to eigh
teen mm long Another tunnel branched o ff f rom the main
channel and going i nto a l ower stratum o f wood there
bra n ched into four sim ilar cell s as shown i n g 2 1 ( sl ightly
enlarged ) each contai ning a de nite success ive stage in the
devel opment o f the young The material used for partitions
w a s the saw dust tightly packed to a th ickness o f ten mm
s o it made a remarkably strong wall
The prey in these cell s w a s two w inged ies o f several

species Two cells that w e re lled and had very small larvae
had four ies each and another had four ies plus one head
I n some cel ls were many heads wings and tho races o f ies
One coc oon was o f a dark brown colo r and all the d eb ri s
i n the cell had been shoved to one end be fo re sp inning ( g
Two adults eme rged on July 1 and 3
.
90
2
9
H T Townsend ] P h or b i a s p [ F K nab ] P horm ia regin a

M eige m [ C H T Townsend ] P s il op us sipho Say [ F
K n ab ] Luc i li a s p [ C H T T ownsend ] E mp ha n op ter yx
H.T Townsend ] S arco
eum y o thy r oi d es Townsend [ C
.
FI G
f ea s t
22
C el l s
an d
the
of
H y p o cra b r o
in
c oc o on
o n e c el l
p i c o lus h avi n g
s ti r
t he
r em a i n s
of
dex ia s p [ C H T Townsend ] Ac ha eton eura s p [ C H T

Townsend ] and S o lva ( Xy l om yo ) p a l li pes Loew [ F
.
'
Knab ]
S ince th i s spec ies i s el sew hei e known only to use brambles
fo r a nestin g place i t i s i ndeed su rpri s in g that th i s one
should have s o s k i ll f ully adapted the ol d tunnel s in the log
fo r he r use I n the stalks o f course there i s n o late ral
space for ind iv idual variation in the nest ; s o it i s all the
more remarkable that in thi s exceptional nest so much good
adaptation was appl ied by m aking cell s rami fying from
one main bu rrow Furthermore thi s nest was constructed
horizontally while all othe r nests recorded have been in
up right stalks
O n one occas ion an elder tw ig was b roken o ff and ex
a m i n ed fo r twig dwel ling insects
but i t w a s unoccupied
In less than ve m inute s w e passed by th i s spot again and
.
'
F L!
CAT CH I N G WA S P S
93
found a H s ti rpi c ol us boring into the so ft p ith I t had

in this brie f time made a hole one S ixth inch deep An
other was found still a t w ork burrow ing in the p ith at s o
late a date as October 2 5
One ne nest in an elderberry twig was found in Novem
ber The bu rrow in the p ith chamber w a s thi rteen inche s
l o ng and one fourth inch i n diameter The gallery was
not at all points in the center o f the stalk but oscillated
from S ide to s ide P erhaps the wasp in digging i t out had
merely followed the l ine o f least resistance and had chosen
the so ftest spots The partitions were made o f the so ft pith
rmly packed togethe r The thickness o f the partition s and
the length o f the cells we re variable as the following table
shows :
.
P a rt i t io n
C el l
in c he s
in c h
inc h
V in c h
in c h
M i nc h
A inc h
in c h
P a rt i ti o n
i f; i n c h
in c h
in c h
in ch
2,
C el l
Vi
2
nch
The uppe r ve inche s o f the tunnel was o pen and unused

All the s ix coco o ns i n the cell s rested with the heavy end
toward the bottom leaving the thin easily opened c o vering
on the top We do not know however whethe r these wasps
habitually emerge by boring through the side o f the twig
o r by struggl ing through the pith to gai n the top exit In
another series o f cells i n a twi g eight inches l o ng the parti
tions in the uppe r hal f were broken and crumbled as i f the
occupants o f the nest m ight have escaped by that way O f
course i f they emerge by the open uppe r end o f the stalk
the priority o f the lowe r inmates o ffers d ii c ulti es These
n ine adults emerged during our absence f rom April 1 2 to
28
Anothe r neat nest ve ry s im ila r to this i s illustrated
exact s ize i n g 2 3 Thi s had fourteen cell s and eleven
.
94
cocoons The cells varied i n size from 2 to 6 mm and

the coco ons tted n icely into them There was no y deb ri s
remaini ng ; every evidence o f the feeding procl iviti es o f
the tenants was gone
P ackard say s that they ava il themselves o f plants whose
stem has a p ith wh ich they can readily excavate and
2
Cresson found thei r nests i n blackberry stems The P eck
hams found that th is wasp uses the stems o f plants ; i n fact
this i s the famous insect that worked fo r forty tw o con
They
s ec ut i v e hours w ith only a ten minute intermission
found al l tunnels o f th i s species to be from th i rty to
forty centimeters i n length and the completed cells con
Our notes
ta i n e d ies o f four d i ff erent species al l dead
S ho w that the species uses various species o f ies P erhaps
P ackard was m i staken i n stating that the females o f sti r
i
u
p
rovi
s
i
on
the
i
r
nests
w
ith
caterp
illars
aphids
o
l
c
s
p
sp i ders and othe r i nsects
C res son says that the Hymenopteron D i o m orus hatched
f rom the nest o f H s ti rp i c olus
.
H yp o cra b ro
c hry s a rg i n us
Lep [ S A Rohwer ]
.
A log o f wood was lying on the ground and j ust beneath

it a heap o f n e sawdust This cone shaped mound p o inted
to a hol e in the log di rectly above P resently a black and
yell o w H c hrys a rg i n us ew into thi s hol e and rema ined
therein until we lost patience Among the sawdust on the
ground were four ies apparently dead but quite f resh
9 S arc o d ex i a s ter n od o n tis Towns
H
C
S a rc od ex ia s p
[
T Townsend ] and A n thr a x la ter al is S ay [ F Knab ]
There was n o w a y o f determining whether the wasp had
dropped them a s sh e entered the nest and had abandoned
,
'
1
2
G ui d e
t o the S t udy
P s y c he
2 : 1
89
of
8 78
I n s ec t s
58
889
6
9
WA S P S TUD I E S AFI ELD
them o r whether she had le ft them here only as a relay

station i n bringing in the foo d S ince the wasp gave no
prom ise o f coming out we open ed the log and got her
The entrance gallery p ierced the hard wood for one and
a hal f inches It had evidently not been made by thi s wasp
but must have been cut by some beetle du ring the li fe o f
the tree because the wounded sur face was all he a led over
Where thi s previous burr o w had reached the rotten wood
beneath the wasp had continued the gal lery The length
o f this tunnel was seven inches not exactly straight but
oscillating a l ittl e while follow ing the same general d i rec
tion The tunnel cam e to an end and a l ittl e to on e side o f
the terminus was a completed pocket containing f our ies
and
I
o
f
P
u
do
mest
i
c
L
H
T
8
ro
m
c
a
a
C
s
E
2
g
[
3
Townsend ]
Nei the r egg n o r larva was found and we could not tell
upon rudely chopp ing open the log whethe r thi s cell was
connected w ith the gallery or even whethe r it was sealed
We suspect that the wasp was only storing thi s cell
B arth has found thi s wasp nesting in an old l og in com
pany w ith Cr a bro o bscur a s C mo n t a n us and C s ex ma c u
l a tus The adults are nectar feeders as reco rded in the
papers o f B ank s and Robertson All o f these records a p
pear under the generi c name Cra b r o
,
P a ra n o thy reus
c i ng ula ti s
P ack
[S
A Rohwe r ]
.
In a steep sandy bank perhaps fteen feet w ide and

twelve feet h igh nea r the margin o f the lake w a s a fai rly
large colony o f these black and yellow wasps There w a s
but at some
no vegetation on this sandy area ( g
recent time a large p il e o f straw had been dumped and
,
T he
g e n e r ic
Thy rc op us
of
n am es
d e R o h w er
th i s
s p ec i e s
e re
f o rm
e rly
'
Cra b ro
an d
FI G
24
n o thy r e us
T he b a re p o rti o n
c i n g u l a ti s
th e
of
s an d
b an
w a s i n ha b i t e d b y P
ara
burned there Thi s i s mentioned merely to show that the

sm o ke and ames had been unable to put the insects to r o ut
A survey o f a large part o f the shore revealed no othe r
colonies o f these wasps While the community was large
noth ing in the way o f communal l i fe could be discovered
Each wasp conducted its own home independently
The open holes were conspicuous because o f thei r form
While the burrows themselves were round the entrance was
shaped l ike an arched do o rway w ith a at bottom and was
one e ighth inch larger than the inne r hole
The was p s
we re coming and going each b ringing a Dipter o us morsel
under he r abdomen da tting into the hole with it w ithout
stopping at the entrance or leaving it at the doorway until
an examination o f the interior could be made
Several o f the wasps were enlarging t he i r nests The
method was to bite out and loosen the sand and rapidly
kick i t out under the body through the doorway where it
.
9s
rolled down the slope below When b iting out the earth
f rom the s ides o r ce il ing o f the nest they turned around
sometimes quite ups ide down and the abdomen squi rmed
as they forcibly attacked each bite In many cases the nests
were under and sometimes concealed by a l ittle natur al
hood or proj ection o f the e arth ( g 2 5 burrow in the up
per le ft hand corne r )
The nests were found when excavated t o be o f various
S hapes The accompanying illustration w ill give an idea o f
the d ivers ity o f form and thei r general nature ( g 2 5 one
fourth natural s ize )
We have never seen lateral cell s di rectly connected w ith
these channel s but have o ften found from two to eight
i solated cell s nea r by P erhaps they had been made as
branches but when the S hort lateral galleries l ea ding to
them were closed the connection could not be di stinguished
in the sand Whether o r not th i s wasp digs a long burrow
and makes seve ral cells f rom the main gallery we cannot
say w ith certa inty I n nests excavated later the cell s we re
sometimes found to be s o numerous that we coul d only
suspect that other wasp s were resp o ns ible for them The
burrow i n all the nests was about one eighth inch in diam
ete r and the entrance one quarte r inch wide When the
mothe r was found with in w ith some ies at the bottom
sh
o
w
ing
that
the
nest
was
being
p
rovi
si
oned
it
was
seen
)
(
that no perceptible chamber or cell had been excavated but
the completely lled and closed cells which we came upon
in the sand were o f very p retty oval form one fourth b y
one
hal f i nch In one such case we found fourteen ies
form ing a pretty heap all p iled up l ike cord wood one atop
the other all w ith the head laid in the same di rect ion I n
two other cells close together twenty ies each were taken
Others contained from eleven to twenty ies All o f the
ies used by these wasp s were P a rali m na a pp en d i cula t a
:
I oo
obstructions n o two burrows were alike O f course there

i s s o me reason for P h ila n thus and Xy l o c eli a m e ta th ora c i c us
making a tortuous galle ry in a clay bank which i s full o f
rocks roots and rubb ish ; but here thi s extreme variety in
habit makes one th ink that e ither there i s instab il ity o f the
speci es or that s o far as natural selection goes th is point
i s o f no consequ ence to the sp ecies ; hence a xed hab it
would be superu o us We l ike to think o f the staple form
o f nest that all A m moph il a p i c tip en n i s make as being xed
by Natural S election that thi s form o f nest i s in some
Wa y best tted f or the needs o f the specie s but in thi s

group no form has a particular a dvantage ove r the others
so none are elim inated and thu s none i s selected
At no o n ( October 2 ) the mothers were all in thei r holes
This may have been due to the cold autumn nights and
mo rnings
Late r in the a fternoon w hen the s un shone
more di rectly upon the bank and warmed it tw o o f the
wasps were carrying i n ies hugging them under the b ody
i n the usual way
In many cases we have found the p rey at the bottom o f
the burrow where i t was being deposited as it was brought
i n b ut we have been unable to nd out i f the wasp completely
closes th i s b urrow a fter the cell has been lled and then
makes anothe r long burrow for the same purpose o r i f
s he makes and ll s one chamber at the end o f the burrow
closes thi s and extends the burrow to fo rm another and s o
on I t seems imp robabl e that a new channel can be made for
each cell fo r in the two days spent in thi s colony we never
witnessed the s ur f a c e closing o f a burrow Furthermore
several ni shed cel ls were found suspi ciously near to an
open burrow and in one nest op ened one such cell was
j ust at the po int o f a sharp angle o f the burrow which
w ould indicate that they are dropped down from the tunnel
One interesting item about thi s speci es i s the tenacity
'
FLY CAT CH I N G WA S P S
1 01
w ith wh ich they cl ing to the y We have caught them in

a test tube and only w ith the hardest shaking could they
be i nduced to rel inquish their hold At the time when w e
were aeld we never th o ught o f the probab il ity o f the i r
holding the y by the a id o f their sting The gure
d rawn from a p inned specimen w hich w a s placed in the
cyanide j ar w ith the p rey and later S haken loose shows
clearly the pos ition o f the abdomen and sting and reveal s
how it i s possible fo r the y to be carried on the under s ide
o f the body w ith the curved abdomen and sting holding
.
FI G
26
P a ra n o thy re us
c i n g ul a t is
T w ic e
n a tu ra l
si
the p rey l ike a hook Th is detail however needs further

clari cation P arke r nds that a near relative Oxy b elus
u
a
d
r
n
i
o
t
a tus
ente rs the nest w ith the y rmly impaled
q
upon he r sting
The only biological note on thi s specie s i s reco rded fro m
Washington S tate by Kincaid who saw it ying about
clumps o f L y sich i to n lea m ts c ha ti ns e
.
Ent
N ew s
1 1 :
35 8
C H A P TE R I V
THE
E NEM I ES O F
Xy l oc eli a m e ta thorac i c us
T H E P L AN T -L I C E
M ickel [ S A Rohwer ]
.
The pi cture herewith ( g 2 7 ) w ill give the reader an

i dea o f the phys ical feature s o f the s ite o f the only col ony
o f X y l o c eli a m e ta thorac i c us wh ich we have observed Thi s
perpendicula r bank six feet h igh was the S ide o f a gully
wh ich had been washed through a vacant lot by many rai ns
The wall at this point faced di rectly southward where the
s un beat dow n w ith burn ing glare
and n o t a breath o f
breeze could enter the hole walled i n by dirt and tall v eg e
.
ta t i o n
On S eptembe r 3 we came here in quest o f large game but

found in stead a swarm o f these tiny black wasps dancing
in the sun on the face o f the blu ff We were startled to nd
them here i n s o great numbers fo r none had been i n evi
dence when w e had vi sited the place only the week be fore
At that earl ie r date however i n d i gg i ng out a P h ilanthus
burrow in th is same bank we had exhumed a tiny cocoon
from w hich emerged one o f the X m eta thorac i c us the rst
we had eve r seen Th is one inci dent leads us to think t hat
the normal time o f emergence o f these wasps was S eptem
ber 1 and that the enti re population had come forth at this
,
T hi s
a sp
o n ly
h a s b een
w w
1 91 6
M r R oh
S o c Am e r 9 : 349
re c t n a m e f o r D i o d o n tus i n t h e P e m p h r e d o n i n i
.
d e sc r ib ed b y M i c el An n E n t
er
ri t es t ha t Xy l o c e l i a i s t he c o r
r ec en t ly
.
1 02
as
t re a t e d b y
Fox
WA S P S TUDIE S A FI ELD
1 04
however they were disturbed f requently ( eigh t times in o n e

brie f resting period ) by other males eager to mate
P resently we realized that the S ize o f the swarm ( o r
rather colony fo r i t was not un i ed enough to be called a
swarm ) was gradually dimini shing ; occasionally one wasp
woul d alight o n the bank and slip quietly into a tiny hol e s o
small as to be a l m o st imperceptible The holes must have
been o f the ir own making for they were so neat a t for the
insects and o f a very uni f o rm s ize We dug out some o f
them to a depth o f two o r three inches but at th is early date
found n o nest They usually sl ipped into the i r holes w ith
out any con fus ion al ighting on the ve ry spot where the
opening w a s Sometimes they almo st ew into the burrow
Thei r ight was a lazy droning ight po i s i ng and waver
ing to and fro i n the ai r more than dancing I t all seemed
the m ore wonder ful because the wasps burrows and al l
were so tiny ; the adult wasp s we re only a l ittle la rger than
gnats
The f oregoing obs e rvations w e re made between 1 1 and
1 2 o clock ; it w a s 3 p
1 11
when we returned to the place
the next day B y that time the wasps were behaving not
at all as they di d on the rst day but this di fference may
have been due to e ither the hour o r the day O ther wasp s
which ind ulge i n these sun dance s in the morn ing o f thei r
l ives usually dance only i n the morn ing o f the day and then
turn the i r attention to more serious o ccupati ons a fter the
second o r th ird day and i n the a fternoons
S
ee
account
(
o f B omb ia n ub i lip en n i s ) It may well be that these was p s
were al ready in the second o r thi rd day o f the i r exi stence
when we found them
On this day m ost o f them when they approached the
bank wou l d fly into these burrows The apertures we re
temp o rarily cl o sed f rom w ith in and the wasp s spent only a
few s ec o u d s there when they w o uld come o ut and y awa y
,
'
E N E M I E S OF P L A N T -LI CE
19
The holes were q u ite inconspicuous and they were per

I f the open ings
c e i v ed only when the wasps entered them
were as inconspicuous to thei r enemies as they were to us
there w a s l ittle need for thei r closing them
None o f these wasps seemed to be making burrows It
i s possible that these were the holes from which they had
originally emerged We s ee no reason w hy they should not
serve per fectly well as places o f n id i c a t io n as well To be
sure the wasp s may have dug them be fo re they were dis
covered but we Sa w none o f them digging at thi s time
On the second day instead o f dancing or d igging a ll
were busy carrying in p rey w hich cons isted o f aphids
Davi s ] These they carri ed i n
A ph is s e ta ri a e Thos [ J
thei r j aws The wasps clung tightly to the i r booty and even
when taken i n a test tube and v iolently S haken they woul d
not release thei r hold
The bank was o f made or lled i n ground with pieces

o f glass crocke ry Cinders etc strewn thickly through it
so it was impossible at thi s stage to trace out the del icate
burrows to the end ; but some that we followed for a dis
tance showed that they we re usually very cro o ked and grad
ua lly wandered downward in thei r course
From time to time until th e end o f S eptember we vi sited
these tiny black wasps i n the di rt bank While they c o ntin
ued pl enti ful they were by n o mean s s o abundant as they
were on the rst day when they per formed the s un dance
Th i s might have been due to the el imination o f the males
The specimens present later and those ente ring the holes
were all the large r ones p robably females and at no time
di d we again nd the mating behav ior ; hence we suspect that
X m e ta thora c i c us m ates once for all time
Approaching autumn did not seem t o cause thei r industry
to wane
On October 6 many o f them were o ut They
were all females c o ming and g o ing busily at thei r nests
,
1 06
carrying aphi ds as usual

No males w ere p resent and
the s un dance seemed quite forgotten
The dance is no
doubt a S ign o f the i r having j ust emerged so p robably the
date we have recorded fo r that event m ight be accepted as
app roximately the date o f the rst appearance o f thi s
sp ecies
,
28
T h e b ur r o
an d
c ell s
of
Xy l o c e li a m e ta tho ra c i c us E n l a r
.
ed
We succeeded at last in tracing one hole to i ts terminus

The opening and gallery were about three mm in diamete r
The channel w a s tortuous ow ing partly to the rocky condi
tion o f the soil and its total length was l ess than four inches
At the bottom o f the gallery ( g 2 8 ) were four aphids
D oubtless the mother was carrying in p rey when w e in
t rud ed
A sh o rt distance t o one s ide in the earth was a
l ittle cell sealed o ff th i s w a s completely packed with forty
eight aphids all o f the one spec i es A ph is s e ta r ia e They
.
1 08
Ashmead says D tristis D a hlb and D m i n uta s Fab bur

row in sand while D a me r i c a n us P ack burrows in hard clay
and makes burrows o f cons iderabl e depth

On D a mer i
c a n us
one o f the tiniest o f all the wasps the P eckhams
have given us a very interesting account and the only rec
o rd extant hereto fore o f an American species o f D i o
d on tus
They nd for this species that in most cases the
aph ids are kil led by squeez ing the neck repeatedly be tween
the mandibles ; in othe r cases the di sturbance to the p rey
i s so sl ight that they are abl e to walk about a s s o on as re
leased The wasp never uses the sting The nests are in
the ground w ith some grains o f dirt i rregularly heaped
a round the edges I t takes the wasp about an hour to dig
the nest ; s he carries the earth out i n her mandibles and front
legs backing from the hole The nest i s not closed until the
p rov i s i on i ng i s com pleted The number o f aphids i n each
o f s ix nests var ied from ve to fo rty and sometimes the
aphi s served a s food for the mother also
4
C era top ho rus
ten or
Fox [ S A Rohwer ]
.
At Valley P ark M i ssouri on June 2 6 we broke o ff an

elder stem and found th is wasp i n the hollow w ithin The
lower po rtion was crammed w ith twenty ve aphids M a cro
s ip hum ru d b ec hi ae Fitch [ J J Dav is ]
The partiti o ns were
o f the pulveriz ed stem I t was 4 a m when dawn was j ust
breaking and w e suspect this mother had spent the night
asleep in he r nest Another elder stem taken near by had
a few loose a p hids o f the same species i n the top cells and
two pupal cases beneath On July 2 0 two a dult C te n a r
emerged f r o m these
,
P s y che 7 : 46
1 89 6
ul l W i s c G e o l
N at Hist
.
B
Com m o
n ly
k w
no
as
S urv
P e m p hr e d o n
2 :
99
1 06
CH A P TER V
T H E B EE
KI
LL I N G W ASP S
P h il a n thus p un c ta tus Say [ S A Rohwer ]

.
The P h ila n thus p un c ta tus i s a f aith f ul l ittle creature but

her task o f nesting i s so p rolonged that we have never been
able to watch the enti re p roce ss in a s ingle case B y putting
togethe r the parts o f the st o ry as we have gathered them we
may be able to get s ome idea o f thi s shy wasp s way o f
l iving
We discovered one burrow al ready begun August I at
8 o cl o ck
A lot o f loose di rt was roll ing down a tiny em
b a n k m en t by the roads ide ; the di rt was f reshly kicked or
pushed out and we ascertained w ith a p r o be that the bur
row underneath i t went into the bank horizontally for ab o ut
one inch We could not rema in w ith the wasp then but the
next morning we were pleased to nd that she was still at
work A good deal m o re di rt had been pushed out We
knew that th is must have b een d o ne ve ry recently because
heavy rains during th e night had tamped down the previous
lot That even ing too when we passed by we were ce rtain
that s he was still at work
Know ing that it would be impossible to watch her fur
the r and desi ring to know whethe r she spent the nights in
he r burrow we dug he r up The hole went hor i zon tally into
the embankment an inch below the sur face level o f the
ground above and towa rd the south for two inches ; then
,
1 09
1 10
it turned at right angles and went westward and downward

at an angle o f forty v e degrees for ten inches thecourse
o f the channel being broken at one po int by a rathe r sharp
kink without any obstruction or cause that w e could s ee
It then tu rn ed at righ t angle s agai n and went di rectl y down
into the earth for two inches H ere at the end o f the tunnel
was the female P p un c ta tus p robably ready to spend the
night The monoton ous detail s o f the channel are men
t i o n ed he re because it certa inly was the crookedest one we
ever sa w i n the work o f wasps N o larva nor prov isions
were p resent the re di d not even seem to be a term inal
chamber s o we suspect that the burrow w a s still in course

o f construction I f th is i s only a part o f her work what
must be the extent o f he r n i shed proj ect ! The entire
length o f the burrow was abou t fourteen i nches and a fter
its many convolutions it term inated about eleven inches be
low the sur face o f the ground
The burrow was already begun when we found it and it
seeme d still to be incom p lete ; neverthele ss we know that
the insect was at w o rk digging fo r two whole days The
wasp is comparatively small and when we th ink o f the small
burrow s o f some o f her l arge r cousins w e marvel at the
amount o f work that s he does I t i s truly wonder ful that
thi s l ittle creatur e should delve so deep into the earth to nd
sa fety fo r her o ffspring which d oubtless she w ill never see
S i nce the top o f the hole i s always cove red w ith the loo se
di rt and si nce she i s neve r seen out o f doors w e suspect

that s he keeps push ing the di rt upwards w ith her head and
out at the o ri ce thereby keep ing her house constantly
closed to intruders
Late r i n the summer S eptember 1 9 we found one insect
hove ring over he r nest which was temporarily clos ed S he
seemed to be per form ing he r ight o f o rientation calmly
po i s i ng i n the ai r and del iberately swinging pendulum l ike
-
1 12
the hole behind her B ut at thi s interesting climax the boys

came romp ing across the diamond ready fo r thei r game so
w e were obliged to relinqu ish the eld to them
Fo r about seven days we had kept our eyes on another
burrow and had known that activ ities were going on i n
s ide i t and a few ti mes w e had caught sight o f the femal e
o r had seen some fresh earth thrown out On S eptember
2 4 we opened the hole and found its enti re length t o be about
twelve inches I t followed a somewhat i rregular slop e o f
appr o ximately th i rty degrees s o the end was ve inches
beneath the sur face o f the ground The mothe r w a s in the
burrow Near the te rminus w a s a S hort branch gallery o r
neck leadin g o ff at a right angle from the ma in channel ;
thi s led to a chamber wh ich was oval and a trie mor e than
a hal f inch long Th i s neck w a s snugly lled w ith so il and
the chambe r contained a white pupal case and some heads
and w ings o f bee s Another burrow o f an identical general
plan a ltho ugh w ith the main channel a l ittl e more cr o oked
n thus digs
leads
u
s
to
think
that
the
mother
P
h
i
l
a
( g
a long main burrow then makes near the end o f thi s a
b ranch w i th a poc ket which she p rovi sions and seals and
that S he then p roceeds to extend the mai n burrow further
w ith a V ie w to making more cell s on the S ides I f this
sp ecies uses one burrow for seve ral young s he i s mo re eco
which p robably digs a new burrow
n o m i c a l than B em b ix
for each larva
Anothe r o f these nests was discovered on July 2 3 I t
p ierced the s ide wall o f a little depression and a large quan
tity o f l oose earth lay scattered beneath it
One o f us
chanced to be near when the P h il a n thus came to he r hole at
a m b rushed out the loose di rt w ith her forel egs
kicking it backwards under her body i n a way very s imilar
to B emb ix and darted into the hol e
Once ins ide s he
.
'
'
BI
E K ILL I N G WA S P S
-
I l
e
o
w
u
m
S
o
o
m
5
9
8
m
a.
9
:
x
o
m
H
o
C
.
Q
.
N
.
1 1
promptly pushed up enough so il f rom the tunnel to close the

burrow Thi s w a s not roughly ki cked out but probably
pushed up with her head since it s eeem ed to b e fo rced out in
cylindri cal form l ike a pl ug At
we agai n V i sited the
spo t and found the hole closed ; but p robably only t em p o r
fo r a probe could easily be inserted The wasp re
a r ily
turned and entered in precisely the same manner as be fore
and immediately closed her doo r aga in from w ithin S he
was not a moment too soon for a red bellied paras ite was
al ready digging in the loos e sand at her heels
Three days late r w e cam e back at twilight and then found
it apparently rmly closed so we opened it Only the upper
few inches were packed w i th soil ; bel ow th is the gallery
remained open I t ran i nto th e tiny bank w ith only a ve ry
sl ight downward slop e for four inches a fter wh ich i t
dropped at a steepe r incl ine about fo rty ve degrees for
eleven inches and then terminated in a chamber an inch
long which turned sharply to th e right The larva in the
chambe r was al ready about hal f grown i t was surrounded
by some deb ri s heads and legs but i n the grow i ng dark
ness we coul d not dete rm ine what the fo o d o f the larva had
been
Thus they f requently i f not usually choose for thei r
nesting site some l ittl e slope or th e s ide o f a depression
where they can dig int o the s ide wall and where the dirt
cast out can roll down the slop e and be out o f the way A
numbe r o f burrows were fou nd go ing di rectly horizontally
i nto clay banks
We chanced to see a S leepy P p un c ta tus j ust getting up
one June morn ing at
S he slowly and care fully
opened the entrance to the hole from within not by push ing
the di rt out as s he does on other occas ions but by ve ry care
fully and slowly working it back into the burrow p robably
by scrap ing it under her Fi rst a l ittl e opening no larger
.
1 1
WA S P S TUDI E S A FI ELD
thus did exactly th e same but when nally

,
he returned to
her hol e w ith a bee o f her own s he seemed to become con

fused a t th e s ight o f ours lying there hes itated and dropped
her own and ew away
The P eckhams have some very interesting notes on the
stingi ng hab its and general behavior o f this was p
They
also illustrate a n es t which i n shape i s unl ike anything we
describe here They nd that the males o f thi s S pecies
construct lodgings i n the sand and return to them night
a fte r night to sleep
,
P s euda n thop hi l us
v er tila b r i s
Fab [ 8 A Rohwer ]
.
O n the a fternoon o f September 1 6 a l ittle p ile o f dirt

about two inches in d iamete r appeared to have be en newly
excavated and lay covering the mouth o f a hole A P ver
til a b ri s alighted on the p ile carrying an insect snug beneath
her body and walked about as i f seeking a place to ente r
She dropped her prey fo r a few seconds while sh e explored
the mound ; thi s gave us the opportunity to see that it was
a l ittle bee all yellow w ith pollen S he soo n p i cked up he r
property again and ca rrying it betwee n he r legs to a precise
spo t on the oppo site si de o f the pile began excavating and
kicking the di rt up beh ind her all the while keep ing a rm
hol d upon her p reci ous bee w ith he r middle legs while the
f o relegs dug up the soil and an instant later the hind legs
ung i t back H er method o f p roceeding w ith her work
seemed to be : a fter S he had gone inside the channel to leave
he r i nsect in the part o f the galle ry al ready cleared retreat
to the opening and kick out the di rt go back and excavate
some more and move the in sect down to the n e w cl ea r spot
,
T hi s
asp
i s g e n e ra lly
k w
no
as
P hil
a n t hu s
v en ti la b r is o r
ert i
B E E K ILL I N G
-
WAS P S
1 1
A fter a m inute or two o f thi s wo rk s he came out and began

vigorously kicking i n the soil At thi s po int we had to take
her o r run the risk o f los ing her i dentity Upon opening the
burrow we found the top tw o inches lled in loosely and the
remainder empty The hol eS loped dow nward at an angle o f
forty v e degrees rst toward the southeast fo r ten inches
then it turned west and then reversed S harply and went di
We could not accurately follow the sharp
re c tly east
curves o f the channel but at the end were found a pupa l case
and a larva both s u rrounded by bits o f black chitin I t was
one o f the longest wasp tunnel s that w e have seen I t w a s
l ittl e wonder that the abundant soil on the sur face made a
consp icuous m o und I s it then the custom o f these wasps to
?
continue using the s ame hol e fo r several o ffspring
I t cer
t a in ly appears s o S ince this burrow al ready contained two
babes o f di fferent ages We do not know whether the bee
which the mothe r wasp was bringing in w a s intended fo r
food for the larva al ready in the nest o r to be a host o f
anothe r egg
The P eckhams have a short note on P v er tila b ri s whe rein
they tell how she takes bees o f seve ral genera and species
into a ground nest S he carrie s he r p rey with her second
pai r o f legs and closes the door wheneve r she leaves the nest
Robertson nd s the adults feeding on the owe rs o f
various species o f S ol i d ag o P ackard gures this species in
the American Natural i st I 77 1 8 6 8 and his G uide p 1 5 8
1 88 5 where he menti ons thi s as

our most common south
ward form
The P eckhams tell us how P pu n c ta tus males make holes
in the sand where in to spend the n ight and we found a
mal e o f th i s speci es P v er tila b ri s h iding one evening at
deep twil ight in the burrow o f a c in c in dela beetle
,
C HA P TE R VI
S O M E M U D D A U B I N G W ASP S T HA T H U NT S P I D E R S
-
S celiphron ( P el op o eus )
ca em en ta ri um
Drury
I n a f ormer study o f the contents o f the nests o f th is

species w e include d many i tems on the behavior o f thi s
insect We p resent here a few notes on the biology o f thi s
species not included in those pages
We learned i n the p revious observations that the mother
sometimes fai ls to supply su fci ent food for the growth o f
the young but w e did not inqui re whethe r the larvae could
ass imilate more than the no rmal amount o f food suppl i ed by
the mother The numbe r o f spiders m ost f requently suppl ied
by her i s s ix o r seven I n the one case in wh ich w e ex p eri
m en ted upon thi s po int w e added three more sp ide rs ; these
were p romptly devoured s o w e added one more B y the
next evening thi s too was gone so we gave the young was p
four more fat ones Two o f these we re eaten and the larva
was found dead Whether its gluttony p roduced its death
w e do not know but i n the ve days it had eagerly con
sumed s i x extra spi ders
During the latter part o f June we had the good f o rtune
to d iscover three P el op o eus larvae i n the act o f pupating
Thi s enabled us to learn accurately the durati on o f the pupal
period at that time o f year One o f them emerged n ine
.
o urn
hav i o r 5
B eh a v i o r
An im al
: 2 4 0 - 2 49
91 5
: 2 7-6 3
1 18
916
Al s o
o urn
An i m a l B e
WA S P S TU DI E S A FI ELD
-
co eruleus that l ived mo re than twelve hours a fter i t was

-
beheaded
FI G
l eu m
30
N e s ts
I n te r i o r
of
c cl i
p lzir o n
an d e x t e r i o r Vi e
5
c a e m e n ta r i um
s
T w o t hi r d s
-
or
Cha ly b io n
n a t ura l
si
ze
aeru
One o f ten p ictures the male wasps o f thi s species as lazy

good for nothing fellows while the females are incessantly
busy bringing in mud and sp iders and mak ing the nest ; but
,
MUD DAU B I N G WAS P S
121
the females to o l ike to loa f sometimes O n S eptember 1 6

1 9 1 4 we observed one mother at work during the a fternoon
the cell was completed but unsealed and unlled
At 3
and the wasp was b ringing mud to rein force the nest by
spreading it thin all ove r the outs ide Up to 7 o clock she
was still thus occup ied w ithout paying any further attention
to the unlled cell Until noon the next day the wasp was
absent possibly loa f ing At 4 o clock when we returned to
the nest we found that the cell had been lled and sealed in
the interval but further w o rk for the day had been discon
We watched fo r her the remainde r o f that a fter
t in ued
noon and all the next day but there was n o return whatso
eve r to the nest On the following day al so we lay in wait
fo r he r and were about to give her up as dead lost or
stolen when s he returned for an hour at m idday and n
i s hed another cell a fter which s he seemed to conside r her
sel f deserving o f anothe r a fternoon o ff Un fortunately we
could not watch her beyond this time but w e have o ften
wondered i f thi s w a s an unusually indolent ind ividual or i f
such i s about the usual rate o f progres s o f the work o f
these wasps
The nests o f thi s mud dauber and those o f Cha ly b i o n
co eruleum are identical Fig ure 30 i llustrates the exterior
and interio r o f thei r homes Some nests S how n icely the de
tails o f architecture ( upper gure ) while in others these de
tail s are obl iterated by the deco rations o f mud pellets ( see
middle gure )
A S ix celled nest o f a P c a em en tari um was found hanging
rmly to a root only two mm in diamete r wh ich ran along its
longitudinal axis I t reminded us o f the mod e o f attach
ment o f the potte r wasp s cell to the twigs o f plants This
was the ro o t o f an ove rturned tree o n the top o f a sand
bank ; the soil was washed away fr o m the roots and on one
o f these strands o f exposed ro o tlet this nest was very de ftly
,
1 22
attached S ince this ro o t w a s s o small it must have t e

qui red a great deal o f de ftness on the part o f the mother
to attach he r rst cel l to i t ; o f course w ith thi s o ne well
ancho red for a foundation i t was easy to build the others
B ut i f this sort o f s ite w a s generally used by p rim itive
a n d thi s undou btedly must have been n ec e s

c a em en ta ri um
sary be fore peopl e provided them w ith barns and shed
o
f
s
then w e think in thos e days o f p ionee ring a deal
ro
mo re o f ingenuity was necessary than i n the l i fe o f ease and
conven ience o f today Campers had built re s in the hol e
under the roots o f the tree and the smoke had blackened the
nest but th is di d not a ffect the contents o f the cells
On another occasion we found a four celle d nest hanging
to a thread l ike v ine wh ich grew int o the shed through a
crack in the wall Desp ite the fact that there was hundreds
o f feet o f board sur face upon wh ich this wasp could have
built i t chose th i s f ra il v in e These facts togethe r w ith
the fact that these nests have o ften been found plaste red to
nai ls protruding f rom the walls on umbrel la ribs corn
husks and othe r unl ikely places suggest that p robably the
p rim itive instinct was to build i n such places
At Lake V iew Kansas a ne large ne st w a s be ing bu ilt
I t was structurally com
o n the outs ide o f a schoolhouse
but
the
mother
was
busy
daub
ing
mud
all
over
i
t
to
l
t
e
e
p
strengthen it In taking i t down it became sl ightly broken
so we le ft it
The next day the mothe r had plastered
mud all ove r the broken part s S ome would say : A W i s e
mothe r l ; but in thi s case there was we b el ieve n o wi sdom

or intelligence at all ; s he was daubing mud all ove r the n est
at the time when the accident happened ; hence noth ing i s
more natural than that s he should continue to sp read mud
all ove r the broken places as wel l In our op in ion it w o uld
have shown real intell igence i f s he would o r coul d have
interrupted her momentum o f sp reading mud all ove r the
.
1 24
the specimens taken f rom the Marquesas I sla nds are i den
tical w ith others which he has f rom the Sandw i ch Islands
Ashmead nd s this speci es paras itized by two specie s o f
cuckoo bees C hry s i s c o erula n s Fa hr and C p erp ulc hra Cr
10
9
Robertson and Cockerel l have w ritten on its o w er f re
1 1
i
n
hab
its
C
H
Turner
has
gathered
some
very
n
u
e
t
q
g
i nteresting experimental data on the homing habi ts o f thi s
sp ecies
.
C haly b i on c a eruleum L inne
While breaking open the mud ne sts o f these wasps at

1
2
p
m
on
July
w
e
we
re
startled
by
a
buzz
We
traced
9
the noi se to our very n ger
tips and ther e in on e o f the cells
w e spied the insect s j aw s p rotruding f rom the pupal sack ;
the young wasp w a s furiously bi ting an opening and all the
wh ile sp itting much sal iva about the po int o f attack I n
less than ve m inutes the new edgl ing w a s f ree and ying
about the room in ci rcles W e wonder i f they sometimes
normal ly emerge at night or whether ou r rough handling
ca used the wasp to make its exit at thi s unexpected hour
I t woul d b e interesting to know too i f the abundant sal iva
i s o f use in breaking down the heavy mud seal o f the nest
H oward in the I nsect B ook records th e obse rvati o n o f
S chwarz H e found C c a eruleum wa s engaged i n c a p tur

ing a certain kind o f S p ide r which hid itsel f so care fully that
i t w a s most di fcult to nd Instead o f S pending the time in
fruitless searching the wasp woul d entangle hersel f in the
web o f the sp ider when the latter would immediately dart
out f rom her hiding place thus exposing he rsel f to the
.
T ran s Ac a d S c i S t Lo ui s
1 0 N t re
1 89 3
a u
4 8 : 438
B iol
B ul l
:21
11
22 2
9 08
45 3
89 4
M UD DAU B I N G WAS P S
1 25
wasp who would easily free hersel f f rom the w eb and
chase the spider to its retreat We have never w itnessed th is

method o f almost human cunning and deception but we are
sure that not every wasp could eas il y extricate hersel f from
the web fo r in four instances at d i ff erent times and places
we have removed f rom sp iders webs the dead remains o f
C c a eru leum
We noticed one C c a eruleum ying and buzz ing from place
to place on the barn wall We lost her f o r a mom ent but
when next seen she had a small sp ider in her mouth and had
her abdomen exed to sting it S he al ighted near by and
it seemed she was tu rn ing it over and over in her m o n th per
haps eating it o r adj usting it for stinging S he eluded close
watching so w e coul d not obse rve the detai ls o f he r habits
The season fo r C c a eruleum is long The extreme date s
on which w e have thus far observed the m o ut were May 2 0
and Octobe r 1 3 They were seen in July and Au gust feed
ing on the owers o f P i mp en ella
A m o re complete account o f the work and behavi o r o f
thi s was p may be found i n our paper on The B i o l o gy o f
Mud daubing Wasps
w ith a description o f the c ocoon
made by the larva rec o rded i n a separate note
Figure 30
illustrates the interi o r and exterio r o f thei r nests
These wasps while they are solitary in the ir mode o f l i fe
by day are gregari ous at night and congregate night a fter
night at a chosen spot for the purpose o f commingling in
sleep The full account o f thi s strange habit may be found
in our pape r on The Sleep o f Insec ts

,
12
1 3
12
1 3
1 4
o u rn
P s y c he
An im a l
2
2
:
6
2
B eha vi o r
63
191
An n E n t S o c Am er 9
.
: 2 7 - 63
9 16
: 2 2 7 -2 7 5
19 1
C HA P TE R VI I
S O M E W ASP S T HAT
P R E!
UPON
C erc eri s f um ip enn i s S ay [ 5 A Rohwe r ]

.
B EET L ES
A Cerc eris f umip en n is at Lake V iew Kansas had chosen
a hard packed highway for her nesting s ite The hol e went
straight down for about two inches and then turned at an
angle H ow fa r it continued w e do not know fo r the road
There was a
w a s s o hard that a trowel could not p ie rce it
The wasp
quantity o f loose soil about the opening ( g
was evi dently digging the cell ; s he went i n to dig for a l ittle
while came out head rst and dropped her loa d at the same
time turn ing ar o und to go i n again and s o continued When
w e crowded to o close to s ee the details o f her work she
would pause in emerging and hold her yel low face aga inst
the aperture as s he s olemnly eyed us for a moment then
slowly retreat and cautiou sly remain w i thin for a long time
,
FI G
31
The m o uth
f the b urr o
1 26
of
Cer cer is
f umip en n i s
1 28
bes ide he r almost touch ing he r lay anoth er beetle o f the

sam e kind the s ixth one which s he had j ust brought in
and dropped The s ite was s o thoroughly mutilated w ith
n o entrance V isible that i t was wonder ful that S he could
recogn i ze the spot at all and f a r more so that s he should
pe rsist in burrow ing i nt o the dishevelled earth i n an a t
tempt to reach the chambe r The appearanc e o f the place
was completely alte red and yet she un falteringly delved into
the heap o f fresh earth at p recisely the right spot to reach
he r old nest H ence unless the sense o f smell was a factor
it was in deed mysterious how s he could w ithout any re
c o n n o i t e ri n g
l o cate the p reci se s p ot in th is large barren
a rea
To S how how ind ivi dual temperament enters into the hom
ing p roblem and how one cannot lay down hard and fast
rules fo r the specie s as a whol e by observ ing a small number
1
o f cases let us cite the interesti ng notes by H artman
on the behav ior o f thi s wasp H e found one carrying a
weevil ( C o n o tra c helus n e ocr a teg i ) into her nest which
was s ituated on the edge o f a v e foot embankment under
a bush about two feet h igh The next day he returned and
cut the bush o ff at the roots and place d it three feet to
the right S oon f um ip en n is returned and ew not to her nest
but to the bush which had been placed to one side

A fter
di scove ring her m istake s he ew away to get anothe r sta rt
came down again from between tw o trees and ew to the
bush S ince she repeated thi s per formance at least a dozen
times w ithout nding the nest it i s sa fe to conclude that it
was the bush that d irected he r ight Moreove r the wasp
always ew down from the same d i rection showing that
earl ie r in her course s he was d i rected by other obj ects es
i
ll
e
c
a
trees
p
y
,
B ul l U n iv
.
T ex
N o 65
.
66
90 5
WAS P S T HAT
P REY O N B EETLE S
1 29
The s ix beetles taken f rom our nest were watched over

care fully but no signs o f l i fe were vi sible w ith all the usual
methods o f stimulation The conclusion was that C fum i
pe n n is kill s her prey outright although it might be poss ible
that these beetles feign death when stimulated
The P eckhams according to Wheeler nd that other
species like the maj ority o f European C erc eris prey upon
weevils but that C f um ip en n is p reys upon a B upresti d
beetle Chrys o b o thris 4 impressa wh ich it kills outri ght

wh ile Ashmead citing H ubbard says they p rov i sion thei r
4
cells w ith C hry s o b o thris d eu tip es and G ro s sb ec k nds that
they use three species o f B uprestids for prey : D i c erc a p n uc
tul a t a B upr est is li ne a t a and Chrys o b o thris ori c ola H e
also says that their mounds resemble conical ant hills which
are abo ut two and one hal f inches in diameter the depth
four and one hal f to six inches and there is no widened cell
at the term inus H i s statement h oweve r that at the bottom
o f some burrows paralyzed B upresti ds were found S hows
that i t is still an unsettled p roblem whether C f um ip en n is
paralyzes he r prey or ki ll s it outright
,
S colia dub ia S ay [ 8 A Rohwer ]

.
While w e had be en exam ining H oward s H ill several times

a week all summer w e had neglected t o penetrate on e small
corne r near the stable s He re w e di scovered S colia dub ia on
S eptember 1 5 1 9 1 5 Whether these wasps had come here
late in the season or whether they had been there for some
weeks i s not known but thi s much is certain : i f they had
3
4
5
n im al
A
n
o ur
P sy c he 7
0 urn
A l a rg e
: 60
.
t rac t
B ehav io
1
89 6
E nt
of
3 : 38 3
913
S oc
la n d
20 : 1
s i t uat e d
35
1912
i n the
s o uth
e s tern
p a rt
of
S t Lo ui s
.
9
3
been there for any cons ide rable length o f time they had
kept very close to thei r own particula r hab itat where w e
found them for all othe r parts o f the el d had been thor
oughly scanned repeatedly Further observations w i l l be
necessary to S how whether they disseminate over the el d
o r keep strictly to a l imited area o f pe culiar character i sti cs
A strip o f ground by the roadside perhaps ve feet w ide
som e twenty fee t f rom the barn w a s thickly overgr ow n
with p igweed smartweed and bulrushes
Th i s ground
sloped down into a depress ion which at that time w a s a
stagnant mudhol e At the end o f th is strip was an old
manure p il e covering a large area partly overgrown with
vegetation Just above these plants and al ong the ro ads ide
a number o f these wasp s were y i ng They were doing
nothing i n particular that w e could s ee but ying idly to and
f ro along th is strip o f ground The i r ight foll owed a
z ig zag cou rse i n and out ; they s eldo m rested and i f one
tried to follow them w ith the eye they seemed to disappear
among the vegetation but one could n ot trace them The
wasps are very consp icuous in ight ; thei r reddi sh abdom ens
w ith the two blotches o f yellow o n one o f the se g ments make
a b rill iant showing There were p robably th irty or more o f
them in ight We watched th em for ove r an hour and
sa w only two o f them s ippi ng the necta r o f the owers o f
smartweed and p igweed and o n e the snakeroot but to
them the sunowers near by d id not seem at all attractive
Thi s strange ight could not have been a search for food
and we hardly think i t could have been a prel iminary o f mat
ing since they di d not seem t o chase one another although
both sexes we re present
Again and again on other days we watched them i n thi s
vague monot ono us ight over the we eds ; we watched in
various kinds o f weather and at d i fferent hours o f the day
S ometimes there were more sometimes fewer o f them but
,
WA S P S TU DI E S AFI ELD
2
3
and alighted on a lea f and rested as long as we were w ill ing

to watch he r ; ou r will ingness extended to one hour and so
l ikewi se did her repose
As the season advanced the behavio r seemed exactly the
same as previously During August the males and females
commingled in the lazy ight to and f ro over the manure
p iles but they di d not pay any heed to each other At one
time during August w e dug up the manure heaps thor
oughly but then found no b eetle larvae
I n S eptembe r only the females w re p resent ; thus it
seems that in the species 5 dub i a a s in many other insect
species the males both appe ar and disappea r in advance o f
the females B y thi s time some o f the females had scarred
and wo rn w i ngs show i ng that they had p robably been active
fo r a goodly tim e
The maj ority o f them we re l urking about the oldest man
ure heaps that had lain undi sturbed the longest time S ince
in p rev ious experi ence we found the beetle larvae only later
in the s eason we can only concl ude here that thes e 5 dub i a
we re l ingering about these p rom is ing heaps wa iting fo r the
larvae to become mature o r abundant in o rder to oviposit
and that they felt s o secu re i n the favorable prosp ects o f the
s ituation that they woul d not even go forth and seek othe r
places but wa ited
We can nd no l iterature on any b iolog i cal asp ect o f this
insect excepting that Ashmead says : Very l ittle i s known

on the hab its o f our forty fou r species o f S col i idae They
are paras iti c on various scarabae i d larvae mostly o f the
family S carabaeidae
,
P s y c he 7
78
89 6
CH A P TE R VI I I
S O M E W OO D B O R I N G W A S P S
-
C erc eri s n i ti m a Cress [ S A Rohw er ]

.
This l ittle creature seems in her behav ior rather para

d ox i c a l
We have three reco rds made in widely d i fferent
times and places o f nding her digg ing i n san d and entering
holes i n the ground in a very fam il iar way From these
observations w e had j ust arr i ved at the deduction that S he i s
a ground burrowe r when w e found one in the p ith chambe r
o f a weed stalk Thi s w a s open at the top where it w a s
broken o ff and s he w a s about four inches down P robably
s he w a s preparing to make anothe r cell
s ince one w a s a l
ready completed Thi s rst one was compo sed o f a layer o f
mud next to the p ith wall a ch amber one th ird inch in
l ength contain ing a pupal case and another layer o f mud
f or a clos ing wall Only ten days later S eptember 9 this
p retty wasp emerged from thi s cocoon the reby proving
that the mother had been there by no mere accident Desp ite
the fact that the stem containing the pupa was already spl it
open on one s ide the wasp bored a neat hole on the other
s ide by which to gain its exit
S ince some species o f Cer c eris use burrowing bees for
prey w emay take the l iberty o f su specting that S he w a s
foraging where w e f o und he r peep ing into holes in the
ground
The adults are necta r feeders d e Robertson and S mith
,
33
WA S P
34
S T U D I E S A F I EL D
tr i dent a tum
Tryp oxy l on
P ack
S
[
A Rohwe r ]
.
A burrow in the p ith chamber o f a ce rtain elder twi g

at Lake V i ew Kansas must surely h ave been an attractive
resort although inconsp icuous to us for i t had evidently

been used tw ice a s the cozy dom icil e o f insects
The burro w i n the tw ig was s ix a n d one hal f inches in
total length I t had evidently been made and used by a n
othe r insect p revi ous to the p resent one fo r the lower three
and a h al f i nches was tightly packed with d bri s o f the
fo rmer tenant At that po int a mud plug w a s rmly tted in
A l ittle way above thi s was another
t o wall up this r e fuse
mud p a rt it io n m a k in g a cell three fourth s inch long and a
thi rd plug was inserted to make another such room There
may formerly have been more cells w e do not know fo r a

mow i ng machine had cut the stalk o ff smoothly j ust above
thi s point leaving the gallery open The neat l ittl e cell s
conta i ned tiny cocoons I t w a s brought in f rom the eld on
August 1 and on August 1 5 an adult T tri dent a tum
emerged
Although thi s species was described in 1 8 6 7 no l ite r
ature could be f o und on the b iology o f thi s wasp
-
'
S i la on
n ig er
R ob
S
[
A Rohwer ]
.
An elder stem w i th a l ittle hol e i n the top aroused our

curiosity O n openin g i t w e found in the p ith chamber
some small pellets o f mud a few b its o f dried leaves thrown
in p romi scuously and then long partitions w ith four c o
coons at i nte rval s among th e deb ri s o f dried l eaves l ittle
p ieces o f mud small stone s and l ittl e pellets o f excrement
-
in
T hi s b e l o n g s
F x
o
s an d
t o t he
W i lli a m s
f a m il y
La rr id a e
a rti c l es ,
an d
de R ohw er
i s th e
.
s am e a s
N i teli op s is
6
3
WAS P STUDI E S AFI ELD
he re illustrate the interio r ( g 32 ) o f an ab andoned cel l o f

the mud dauber partitioned and res c al ed and containing two
clav a tum coc oons w e have al s o long suspected that they
adapt and use in the same manner the holes al ready made by
othe r insects in logs and fence posts Knowing f rom our
observation s that S he f requently makes her nest i n ol d
borrowed cells our surprise was great when w e found that
T cl a v a tum can al so de ftly tunnel into wood and make her
own burrow This S how s a greate r d iversity in activ ity than
is o ften found in wasps I t i s indeed wonder ful that so
high a degree o f special i zation S hould be associated w ith
such divers ity o f accompli shments o r habits as we nd in
th i s wasp
.
FI G
s ui t
si
ze
32
t he
T he
n ee d s
a b a n d on e d
of
c el l o f
Try p ox y l o n
m ud - d a ub i n
l a va tum
P up a l
gw
a sp
ca ses
in
rem o d e l l e d
s i tu
to
Na t ura l
They work when burrow i ng in a ve ry cha racteristic and

mechanical manner
O f course w e could not s ee the ir
method o f cutting the wood but they get thei r load o f s a w
dust in the depths o f the burrow emerge backwards and y
away backwards a few inches fro m the mouth o f the hole
hover or poi se on the w ing for some seconds until they
d rop the load o fttimes go ing mechanically straight to the
hol e again and sometimes turning th e body S l o wly around
and ying in a circle wh ich o f course makes a very stiff and
ungrace ful movement and then returning to the hole
A number o f these l ittle wasps were found l iving in a
.
WOOD
B O RI N G
WA S P S
37
sem i gregarious way in a wood lot While they were fairly

numerous about the hal f dozen long rows o f cord wood
there w a s only o n e sp ot at the end o f one row where from
ve to eight indiv iduals congregated at a time for play They
would y about the top o f the p il e o f wood at a height o f
ve feet chas ing one another on the w ing for an hour or s o
at a time do ing noth ing in particular but to play thus in the
B oth sexes were present s o we thought it m ight pos
s un
s ib ly be a coquettish behavior p reparatory to mating
A
certain scar on one o f the logs where a branch had been
cut o ff w a s the one and only place where they would rest
One at a time they would select th is spo t ignoring all the
s im ilar ones at hand and remain there quietly unti l di s
covered and put to rout by the othe rs The latte r would
chase her ( i t w a s usually the females who sa t d o wn to rest )
back into the crowd During the a fternoon we sa w ve
group s there i n turns frol icking i n the s un ; but the beha
v io r never var i ed the location never changed and the cut
end o f thi s one log never se emed to g row less attractive
Yet during all thi s pe r formance w e saw no attempt at
mating
-
Try p oxy l on
o hns on i
Fox [ S A Rohwe r ]
.
Fo r o ur knowledge o f this S pec i es we must depend largely

u p o n me re surm ise or regula r detective methods We have
seen it active only once when at Lake V iew Kansas on
July 2 7 it entered three hol es in a clay embankment What
its errand there w a s w e di d not ascertain whether searching
for a domicil e or hunting prey
A year later we got our next c lii e to he r ways o f l iving
A stem o f some s o ft wood was found near to the ground %3
inch in diamete r and 1 3 i nches long It had been hollowed
-
WA S P
8
3
S T U D I E S A F I EL D
out and contained mud partitions at interval s o f

i nch
3
1
1
and
inch
There
were
also
indi
2
V
A
cation s o f forme r mud partitions which ha d been removed
There were also S ix holes in the stem al ong one S ide These
clearly show ed by thei r large rough and asymmetrical form
that they had not been made by the insect em ef g in g f rom
the i nterio r but must have been made by a stout j awed or
toothed creature f rom the outside These openings w ere
o n the outsi de i rregula r and var i ed f rom
to
inch
i
n
M
length while on the ins i de they were %3 to
inch i n length
We can only suspect that a wood pecker o r some food
hunter o f l ike hab its had broken into the nest when it was
rst used and devou red the occupants and that therea fter
thi s wasp rented the house i n sp ite o f the leaky walls At
the bot tom o f the cavity was a dried mass o f sp iders legs
and parts Whethe r the mother wasp had lled the lower
cell w ith S p ide rs and then forgotten to lay the egg or
whet her in renting and clearing th e house S he had shov ed
back all the d bri s as does T cl av a tum we do not know
S ince there were no shedding sk ins or other rubb i sh w ith
the S p iders i t seems sa fe to say that these sp iders belonged
to the latest o ccupant
Just above thi s compartment full o f spider remains w ere
two cells each contain i ng a goo d pupa wh ich later gave
fo rth adults o f T j ohns oni
N ext above these was a sol i d mud partition o r plug wh i ch
was unbroken ; th is s he had ev i dently put i n fo r he r roo f
One o f the S ide hol es had served a s her doorway I t i s
evident that thi s had been the upper term inus o f her nest
fo r thi s heavy partition was unbroken while all those above
this were pe rfo rated as i f the em erging insects had es
caped by the top o f the stem and the upper cell s were empty
Thus only by v irtue o f detective methods but w ithout
having been a w itness to the deeds w e suspect that thi s
,
CH A P TE R I X
TH E
A ly son
H U NTER S O F S MA LL O R T H O PTE R A
melle n s Say [ S
.
R o hwer ]
The expe r i ence o f the summe r s eld work in di fferent

places has taught u s to nd these delicate creatures only in
the pecul ia r env i ronment that seems best adapte d to the i r
needs We have never found them elsewhere than in a cool
damp bank o f mud o r sandy clay nea r to a b o d v o f water
They may be very nea r to the dry earth or the hot sandy
areas but they scrupulously avoi d such regions
The rst ones that w e found were at Moselle M i ssouri
on J une 30 1 9 1 6 They were seen carrying green insects
unde r the i r abdomens and entering holes in the mud bank
The i r burrows w e re inv i s ible to us because o f a numbe r o f
l ittl e wet p el lets o f mud that l ay over the mouth o f each
hole but the owners found thei r w ay i n somehow A fte r
the wasp s were captured the holes were exposed by brush ing
away the pellets cover i ng them S ometimes the l ittle lumps
o r rathe r shapel ess mouth ful s o f mud were strewn or
heap ed w ithout any a rrangement ove r and ar o und the
mouth o f the burrow ; i n other cases they were piled dex
t ro us ly about the hole to f orm a crude l ittle chimney o r
canopy ove r the o rice on the slop ing bank
Thei r l ittl e burrow s w ere about o n e e ighth i nch i n diam
eter and w ent stra igh t into the bank at right angl es to its
sloping sur face t hen curved S l ightly downwa rd They varied
I 4O
H UNTERS O F S MALL O RTH O P TERA
1
4
from one and one hal f to
in length
four inches ; the f ew that
we f ound completed term inated in a l ittle oval pocket ( s ee
illustration g
which was merely the end o f the chan
nel slightly w idened into oval form
We di d not chance at any time to nd a cell already lled
.
FI G
33
Aly s o n m e lle us he r b urro w

,
an d
rey
N a tura l
Si
w ith p rovi sions at the end o f an open burrow but i n the

earth nearby w e found on m o re than one o ccasion l ittle
cells wh o se connection could not be tra ced These we re the
term inal cel ls o f other n earby nests wh o se channel s had
been closed The rst o f these i solated cell s was packed
tightly with a number o f lea f hoppers There were about
a dozen o f these frail creatures s o delicate that we coul d not
handle them eve n w ith forceps but w ith a straw we gently
brushed them into a vial They exhibited no S igns o f l i fe
and truly we cannot see how such frai l organi sms could sur
vive even the handl ing o f the wasps much less thei r sting
Twenty hours later when we exam ined the hoppers wh ich
we had placed in a sealed V ial every one was shrivelled and
hard s o des icc ated that a del icate l arva could hardly derive
any nourishment from th em This di scovery suggested a
,
42
clue to the p robabl e reason for this wasp pe rs istently choos

ing a damp cool place for he r nest : only in such surround
ings coul d the food be kept fresh and mo ist The wasps con
tin n ed to be se en during all o f the s eason up until October 7
in w idely separated places but always in permanently damp
situations
S ome variation oc curs i n the form o f the top s o f the nests
At M osell e the ente ring wasp wo rked its way between the
pel lets o f a l ittle mou nd o f loose soil and these woul d again
fal l together and cove r the hole a fter the wasp had d i s a p
r
w
ithin
There
were
others
w
ith
a
l
ittle
ch
imney
or
a
d
e
e
p
canopy o f mud At Lak e Vi ew Kansas in the sand a few
feet from the water s edge w e found several nests under
scattered dead leaves i n w h ich case the loos e sand was p iled
behind the hole i n a l i ttle mound and the open ing was i n the
s ide o f thi s mound We even found two n ests at the base
o f a hillside very ste ep at thi s poi nt twelve inches above the
l evel ground H ere the holes went i n ho rizontally at right
angle s to the sur face for a l i ttl e way and then turned
downward M uch loose soil had rolled down the steep i n
cl ine beneath the ori ce and tw o beauti ful canopies made
by the wasp s proj ected for one fourth inch above the
holes We were su rp ri sed to nd A me ll eus i n s o dry a
place as th is was but when w e dug down to the chamber we
found the earth very damp and gummy
The group o f them a t Lake V iew Kansas on a plot o f wet
sand near the the lake shore conducted themselves i n a way
that puzzled us They we re all running thi s way and that
with heads close to the ground as i f anxiously searching
for something They looked mo re l ike a group o f large
red ants in con fusion than wasps All thi s activity was
con ned to a space less than fou r feet S quare and it went on
for over a hal f hou r while we could remain to watch them
yet they di d not nd what they wanted At one time one
,
44
extend downward in a gentle S lope for a distance o f twelve
i nches to a chamber one hal f inch in diameter wh ich con

ta in ed seven lea f hoppers
-
Sm i th [ S A Rohwer ]
Ta c hy s p hex term i n a tus
This energeti c wasp goes about its nest bui lding in a

manner very S imilar to that o f B em b i r n ub i lip en n i s I t digs
i n sandy places ; i t attacks a spot furiously and digs down
rap idly w ith the forel egs inging the dirt out unde r the
body and beh ind it Occasionally it backs out o f the hole
to sweep back the accumulating sand in f ront o f the door
It seems to have l ittle di f culty i n digging i n sand ; the w o rk
goes on qui ckly and somehow i t contr i ves to keep the
r o o f from fall ing in We have watched many B emb ix sp i
n ola e commence holes
work them fo r thre e o r four inches
and then abandon them but not so w ith this species ; whe re
S he starts work S he holds he r Ow n and seld o m changes her
m ind There are exceptions to th is rul e o f cour se We saw
one attempt twelve d i ffe rent holes i n a small a rea whe re our
f ootstep s had al ready lo osened th e sand s o o f course her
roo f woul d not hol d rm ; but as soon as the roo f caved i n
Th is however w a s not
s he would start i n another place
the fault o f he r ca reless w o rk o r her c k len ess
Th ey dig w i th remarkable S peed and skill when work
ing in sand and contrive to make a s imple but shapely bur
row ; but w hen working i n harder soil or clay they dig very
di fferently We p resume thi s di ffe rence in hab it i s due to
The
the d i fference in so il and not at al l to the locality
sand diggers we re not more than forty m iles di stant from
the clay diggers
The latte r wh i ch we observed seemed
j ust as intense in the i r work as those j ust described but
seemed to experience more di f culty in accompl i shing thei r
-
HU NTERS O F S MALL O RTH O P TERA
45
task
They would o ften begin digging several places in
succession be fore they carried a b urro w to its completion
They would usually begin to dig in some l ittle depression or
shallow hole where thei r hard work was already begun for
them H ere they kicked out the loose dirt and then en
la rged the hole su fci ently to rece ive thei r stock o f p ro
vis ions The resulting burrow was not o f any d enite form
or dimensions however but o f any shape and S ize that the
wasp could best excavate in th e hard earth Sometimes her
burrow was barely su fcient to get he r i ll arranged mass o f
p rey under cover She attacked the hard soi l in much the
same manner as she w o uld loo se sand ; hence i t i s not s ur
p rising that S he was not very ei c ien t here Sh e had n o
rened methods for smoothly cutting out the hard earth as
has O dyn erus or A mmoph il a May it not b e that the habit
o f digging became xed whi le the species inhabited sandy
areas and that he r old habits hampered her when S he wan
dered to a d i ff erent envi ronment and attempted to carry on
her work by old method s ?
The burrow when compl eted i s merely a short curving
channel go ing into the sandy earth aslant ; i t i s about three
inches long and the termi nus i s ab o ut an inch and a hal f be
low the sur face o f the ground ( g 34 sl ightly enlarged )
There i s n o real p oc ket or cell These wasps are very wary
about the i ntrusion o f human beings so i t i s diffi cult to
draw near enough actually to observe thei r ways without
frighten ing them away
The nest i s s o on prepared and the wasp ready to begin
carr ying in the O rthopterous prey When at last the nest
i s provisioned S he closes her burrow with l o o se sand in a
way very s imilar to B n ub i lip en n i s ; S he emerges kicking
the d i rt unde r her into the burrow and a fter it i s fully
T he La r ri d a e , t o
w hi
ch
thi s
a sp
b el o n g s
a re
d ig
er s
in
sa n d

closed takes great pa i ns to kick plenty o f lo o se sand over
the hole to conceal i t com pletely When one p roceeds to
dig i t out one nds the sand s o loosely packed i n the
channel that i t can readily b e tickled out w ith a gras sb la de
One mother carrying home her p rey exper i enced some d if
culty i n nd i ng the depress i on where her nest w a s located
,
FI G
34
T he b urro
f T a c hy s p hex ter mi n a tus
S l i htly
en l ar
ed
S he laid down her O rthopteron w h i l e examining spots on the

ground but woul d always p ick i t up when ying about
getting he r bearings ; this she did several t i mes ( s he seemed
a f rai d to leave it lying exposed on the ground while S he
went far away ) carrying i t under her abdomen and ying
sw i ftly and smoot hly as though S he were not s o laden
and once on the way even engaging in a ght w ith a B sp in
o la e
At last S he d ropped her burden in the depression o f her
nest kicked out a few strokes o f di rt entered the hole and
dragged he r hopper i n a fte r her
Fo r some time w e had suspected that they build more
than one nest in a place We had been watching the pro
gress o f a ce rta i n nest f or most o f the day Late i n the
,
48
w ithin Thus S he was ready to spend the ni ght And so

our susp icions were conrmed that T term i n a tus makes a
burrow i n the evening sleeps in i t that night ll s it the next
morn ing and be fo re evening makes another for her next
night s shelte r and so on The problem o f where S h e S leep s
at othe r times than i n the nesting season remains unsolved
I n digging i n anothe r local ity we accidentally ran across
anothe r series o f ve cells one right a fter another and abou t
one inch apart Th i s also lends strength t o our bel ie f that
S he builds a chain o f nests This may account al so for the
fact which we have mentioned be fore that s he makes no
ight o f o rientation ; with so many n ests in one local ity s he
soon becomes s o famil iar w ith the region that S he needs no
special study o f its landmarks
The prey o f thi s wasp i s O rthopterous insects They are
usually o f th e s p e ci es S y r b ul a a dm i ra b ilis U hl [ A N Cau
dell ] small and medium S i ze nymphs In othe r cells how
ever w e f ound one O ed ip o di d nymph [ A N Caudell ] and
o n e small D i c hr o m orp ha v i r i d is S cud
A
N Caudell ]
[
nymph i n the rst stage Ac ri dea n nymphs and nymphs o f
M elan op lus s p [ A N Caudel l ]
From ve to nine i nsects
are usually used to prov ision a cell
Whethe r or not there may be any colo r relation between
the wasp larva and i ts food cannot be declared ab solu tely
but we were i nterested in observing that one young larva
while another on a
on a brown hoppe r w a s itsel f brown
green food was a s green as its host I t has been suggested
that thi s colo r i s p robably due to that o f the j uices o f the
i nsects consumed
S ometimes the larva was found adhering t o its food
insect between the second and third pai rs o f legs on the
ventral S ide ; i n othe r cases they we re on the unde r s ide o f
the neck where i n making thei r attack they had promptly
severed the head f rom the body
.
H U NTERS O F S MALL O RTH O P TERA
49
The hoppers as a rule survive the maltreatment o f the

wasp only feebly Many o f them the maj ority o f the small
ones are killed outright while only the most v igorous sur
vive a day or so and they can respond but very feebly to
stimulation
Ashmead treating o f this species under the name o f
L a rr a term i na t a nd s that they prey up o n C h or top hag a vi r i
dif as c i a ta ; Will iams took one o f these wasps with a young
B anks nds they frequent the owers o f C ea n o
T ry x a l i d
,
thus
Ta c hys p hex
fusc a s
Fox
[ S A Rohwer ]
.
It was late i n the season fo r wasp s to be active yet thi s

little grey T fusc a s was very busy on thi s mo rning o f
Septembe r 30
S he ew to an abandoned building and
alighted in the mortar between the rocks o f the foundation
S he had previously commenced he r hole there ; n o w S he
vigorously kicked back the loose mortar and it fell to the
earth below A fter about ve m inutes o f thi s arduous work
she walked a few feet away and cleaned hersel f Then s he
turned to anothe r s p o t on the ledge o f the foundation and
began to dig in the loo se so il lodged on thi s kicking the ne
earth backwards unde r her body as S he had done in the other
instance One large lump o f something impeded her p ro
gress s o s he snatched it up bodily carri ed i t some distance
i n her mandibles and threw i t away ; she then ew a few
feet sat daw n and cleaned hersel f again and p resently
started to dig anew in the th i rd place A fter only a few
seconds here s he moved agai n and commenced a fourth
hole It seemed that she could succeed in digging into only
the stratum o f loose di rt and when s he struck the har d
,
o
5
cement s he gav e up the proj ect and started a new

S he
wo rked away here for ove r an hour kicking the dirt behind
he r and as the hole deepened occas i onally com ing to the
opening and kicking i t furthe r back out o f the w a y S he
o ften emitted a sharp s queaky buzz whil e s he w a s intent
upon her wo rk i n the hole Two trains pass ing near by
though they made a great uproa r a s they entered a tunnel
unde r the hil l di d n o t di sturb he r i n the least Whe n w e
returned i n the a fternoon w e found that thi s hole to o had
been gi ven up and despite the long t i me and arduou s work
expended upon i t i t w a s only three quarters i nch deep We
s a w her n o mo re
A second wasp o f th i s k i nd w a s d i scovered about n o on
o f the same day j ust commenc i ng her bur row S he chose
a strange spot an ant l ion p it at a point hal f w ay down the
slop i ng S i de o f the funnel and dug i n ho r i zontally A fter
hav i ng w i tnessed the astoni shing pe rs i stence o f the rst wasp
i n wo rk i ng i n the plaster o f the ol d wall w e had supp osed
that some s im ilar s ite w a s absolutely necessary to their
happ iness s o w e were surp rised to nd another us i ng s o
di fferent a kind o f place Evidently T f us cus i s a versatile
l ittl e creature We longed to stay and watch her but tra in
time w as approach i ng and the complet i on o f the story had
to be le ft fo r anothe r t i me
.
Ta c hy tes
o bs c urus
Rohwer ]
Cress and T
.
ob duc tus
F ox
8
[
Th i s smal l T o bscur a s w a s f o und struggl ing w i th a S hort

horned locust S he mounted it and dragged it a few i nches
grasping the hoppe r by i ts antennae S he tr i ed to hol d it
w ith i ts back against the ea rth but had d i ffi cul t i es i n keep i ng
.
2
5
N o tog on i d ea
rg e n t a t a
E ve
[S
A Rohwer ]
The appearance o f N o tog on i d ea a rg e n t a t a early in the

spring has long puzzled us On the morning o f April 7
191 5
we s a w at Cli ff Cav e perhaps two dozen o f them on
the sunny S ide o f the blu ffs apparently do ing noth ing in
particular resting stroll ing or ying about in a sheltered
spot in the sunshine A l ittle later in the day w e found the
same behavio r i n a group i n a sunny southern exposure I t
was surp ri s ing to nd them out s o early and we still wonder
whethe r they had hibernated or had e merged s o early from
thei r origi nal burrows
B ut although they were seen thus in group s i n the rst
d ays o f sp ring they do not continue the gregarious habit
but work s ingly during the summer mont hs
On July 2 5 we fell to watching on e a s she was walking and
running in and out among th e sho rt grass along the r o ads ide
in an open eld S he soon darted into a hole i n a small S lop
i ng bank The burrow was shall o w for w e could s ee her
l ittle black and grey b o dy only a l ittle w a y beyond the aper
ture
As S he worked s he occasionally pushed out the
loose soil beh ind her and it fel l out o f her doo rway and
rolled down the m iniature hi lls i de P resently S he le ft her
work i n the burrow and walk ed to a n d f ro around its O pen
ing ; i t w a s not long be fore w e real i zed that S he was not
makin g he r hole but w a s at work ll ing i t up S he di d not
try to put back the di rt that had been pushe d out because
that had rolled down the S lop e n o r did S he carry it back in
her mandibles I nstead S he p icked up pell ets fragments
o f stems
dried leaves and any availabl e deb ri s that lay
about and dropped them into the hole All o f thes e s he
,
w w
k w
M r R oh
N o tog o n i a b y
i s en e ra l ly n o n h a s b e en c ha n g e d to N o tog on id ea
n a m e h a s b e en u s e d f o r m o l l us s
7
er
r it e s
tha t t he
n am e
w hi
ch
sin ce
thi s
the
fo
a sp
m er
HUNTERS O F S MALL O RTH O P TERA
53
gathered within a radius o f twelve inches from the nest

The great maj ority o f them were taken from e ither the right
o r the le ft s ide o f the hole where she could wo rk on the same
l evel ; a few times S he brought a f ragment from above the
hole but only once or tw ice did S he attempt to bring any
thing from b eneath the burrow Occas ionally s he would
tug away at something to o large or too heavy for her but
S he appeared nervous and dis
s he would soon l et it go
crim inating and seemed to have a hard time in nding j ust
what s he wanted Fearing to lose her and her identity w e
took her when the hole w a s nearly l led
Upon o pening the burrow w e had some di fculty in
accurately following its contour because o f the entanglement
o f grass roots The hole seemed to be about t w o and one
hal f inches deep The lowe r th ird was a chambe r contain ing
three cri ckets o f two species Gry llus p ennsy lv a n i cus
B urm nymp h [ A N Caudell ] and N em o b i us f as c i atu s
D e G [ A N Caudell ]
Abov e thi s the gall ery w a s all
packed w ith so il The wasp must h ave used all the available
di rt fo r the hol e at the time when we arrived up on the scene
Only the upper one thi rd o f the bore w a s lled with d bris
When w e rst opened the chambe r and found the l ively lot
o f crickets w e thought we had made an error and ha d
broken into a crickets nest Soon w e spied the wasp s egg
white and slightly crescent shaped fastened to one o f the
crickets where the leg j o ins the body and curving around the
right side unti l it was vi sible from the dorsal side The
crickets were active when stimulated i n the legs antennae
and mouth parts and while they could not actually hop
they could walk f o r a few steps when p rodded All three
were males I n al l p robability the egg w as depo s ited at
about 2 :30 p m but we could not tell whether i t had been
deposited on the rst sec ond or last cricket b rought in B y
the next morning they had all walked some distance from
.
54
where they had been placed in the j ar ; even the one bea ring
the egg had wandered away All responded actively to stim
B y 7 o clock o f the second morn ing the egg had
ula ti o n
hatched even though its cr icket host w a s stil l pers istently
active The larva was a del icate l ittle yellow creature not
ver y fat I n sp ite o f its small S ize it was a l ittle gourmand
and by the second day o f i ts li fe it had devou red almost all
o f the rst cricket The othe r two crickets were still al ive
and w e wanted to keep them to test the ir longev i ty so w e
substi tuted f o r theni another and la rger S pecimen o f the
same sp eci es We cru shed its head b e fore p lacing it w ith
the larva lest by i ts act i vity it m ight do some inj ury to the
latte r B y 9 o clock that evening the larva had devoured
the contents o f th is one also leav i ng only a thin dry shell
The in fant grew w el l an d seemed to thrive on thi s s o that
night we ga ve it another newly killed cr i cket o f the same
k i n d x Thi s o n e was large r than the others

almost full grown
so lest the integument would be too tough a few i ncis ions

were made i n the abdom inal wall be fore se rving i t The
larva w a s placed thereon and it fel l to at once greed ily de
B y th i s time th e smal ler cr i cket wh ich
vo uri n g the j uices
had constituted the original p rey had grown quite feeble
but the larger one w a s still active
W ith the i ntroduction o f the last cr i cket the larva had
had three the number originally p rov i ded f or i t by i ts
mother ; but tw o o f these we re much la rger than those
which she had suppl ied We did no t presume to inter fere
further w ith the mothe r s p rovi sion by o ffer i ng i t more
food once i t had consumed th is yet it fa i led to pupate and
eventually died a fter th ree days more probably because i ts
nursery w a s not properly equipped or regulated The other
t w o crickets which had be en stun g and buri ed l ived four
da ys
Almost a month late r August 2 2 we were scouring a
.
'
6
5
o f the nest as soon as it w a s opened and even at ten o clock

that n ight all o f them could walk o r hop w ith alacrity upon
S l ight stimulation Judging f rom the fact that the egg was
on the top cr i cket and the wasp w a s clos ing the nest when
,
E
FI G
35
N o tog on i d ea
a rg e n ta ta
her b urro
an d
N a tura l
si e
arrived w e can sa fely estimate that the egg was laid

between 1 and 1 30 p m
On the evening o f the next day the c ri ckets were all as
to n i s hi n g ly l ively ; the one bearing th e egg ran and hopped
clear acros s the table when the box w a s opened A fter
four days i mpri sonment they were a l ittl e subdued and
seldom walked about i n the box B y th e even ing o f the
2 7 th ve and a quarter days a fte r the clos ing o f the nest
the smallest cricket w a s grow ing lethargic and by the 2 9 th
al l but the largest which bo re the egg were dead and i t too
seemed almost gone The egg never hatched
8
Ashmead says under the name o f L a rra a rg e n ta ta that in
the south this insect stores as many a s S l X i mmature crickets
which s he completely paralyzes The w ay the crickets here
j ump out o f the nests when they are opened readily shows
that the M i ssouri wasps very in c o m p l etely paralyze the ir
p rey
F i gure 35 shows anothe r nest i n cou rse o f construct i on
we
P s y c he 7
63
896
OF
H U NTERS
S MALL O RT H O P TERA
57
which w e opened The mother was at the bottom o f the

bur row
Williams gives some interesting data on the biology o f
thi s species H e records that one soon rendered helpless a
small Gry llus by a sting under the thorax malaxated it on
the ventral s ide o f the neck and carried i t towa rd her nest
ventral S ide down The wasp made l ittl e runs and sho rt
ying j umps w ith her burden and a fter s he entered her
hole remained there fo r ve m inutes
I n lling up her
tunnel s he gathe red l ittle lumps o f earth and other material
such as twigs thorns and grasshoppe r excrement and
dropped them in None o f the material he says w as tamped
down She did no t us e he r feet in lling up the hole ; the
feet o f N o tog o n i dea a re not armed w ith brushes to assi st in
such work and the heavy black earth in which s he was
working did not readily perm it digging w ith the feet
In the rst arg ent a t a that we obse rved w e found precisely
thi s behavior ca rrying and d ropping into the hole pellets
fragments o f stones an d leaves Upon open ing the burrow
howeve r w e di d nd the gallery j ust above the chambe r
well packed w i th earth wh ich she surely must have c o m
pressed not in the usual w a y i n which wasp s pack down the
s oi l but as our second wasp showed w ith the ve ntral part
o f the abdomen moving i n a slow grinding c ircular motion
While we di d not witness all the ll ing o f the hole o f the
second wasp w e did see this one scrap ing i n loose dirt w ith
the f ront legs A thi rd one to as we have shown used
he r legs to scrape i n the soil
S ince w e have recorded indiv i duals o f thi s species very
early i n the season one woul d expect that the i r vegetative
and reproductive functions would soon be ove r and they
would die o ff B ut th is i s not s o for we have one record o f
.
Kan s
U n i S c i B ull 8
.
: 1
91
913
s
5
having seen a dozen o f them out foraging on October 6 at

S t Loui s and othe rs as late as October 2 8 at Cl i fton Ter
race Ill ino is evidently st i ll i n quest o f p rey The n i ghts
at the latter time were very cold We wonder i f thi s i nsect
does not h ibernate i n the adult stage ; i f so w e may nd
them p roducing more than one gene rat i on a year a condi

t i on wh ich i s unusual fo r the wasp s of the group t o wh ich
th i s one belongs
We have yet d i scovered no enem i es o f th i s spec i es A fter
a heavy ra i n on August 2 0 however a few o f the wasps
were found dead i n the eld ; they seemed somehow to lack
the means o f protect i on f rom the assaults o f th i s kind o f
weathe r
,
6o
FI G
l eg s
to
36
P ri on o n y x
w
w
il
l g g
atra tum e x p res sly

,
c a p a b l e o f s c ra p i n g a n d c a r ry i n
g so
ra c e u l ly st ra
dd le
a n d r id e
ho m e
d ra
ar
as
to
ell a s
ra s s
S ho
b ei n
ho p p e r
w
gl g
th e
on
on
d er f ul
gh
iz
en o u
N a tu ra l
was about That done s he too k a careless free ight o f

about eight feet looked aro und for only a m inute or two
and began industr i ously digging I n about ve m inutes
s he had dug to a depth o f one thi rd o f the l ength o f her
body ; th en quic k a s a ash s he j umped out and w ith hal f
a dozen rap id scoop s whi pped all the di rt back i nto the
hole and heaped i t neatly on top Then she t ried a th i rd
place l ess than a foot away but dese rted that als o a fte r a
few mouth fuls had been taken out and replaced S he began
the fourth burrow six feet f rom the last a fte r walking
about ten m inutes ; she dug a hole the length o f her body in
a very bus i ness l ike manner then j umped out and kicked
the di rt in again N ext she walked about fo r ten m inutes
taking out a fe w mouth ful s i n othe r locations but putting
the dirt back w ith the same ul tra precisi on as be fo re and
at l ength ew away over the grass and across the eld for
s ome di stance B ut s he seemed fa ith ful t o h er original
purpose ; a fte r ten m inute s or so S he came ying back
al ighted on the bare spot and doggedly began another hole
In seven m inutes it was as deep a s the length o f her body ;
then S he went o ff across the grass leaving i t open
In
about ve minute s she came trudging back w ith a huge
.
H U NTERS O F LAR G E O RTH O P TERA
61
hoppe r ; we do not know whether she had j ust caught it or

had had it there somewhere all the time du ring her many
attempts at digging in this region ; but from what we sa w
later we j udge the wasp undoubtedly had the hopper in hid
ing She carri ed i t fully three feet beyond he r h ole and
hunted fo r ve m inutes be fore s he rediscovered her burrow
S he then resumed digging earnestly going in and out me
c ha n i c a lly l ike a toy w ound up and coming out two or three
times to ki ck the di rt further back from the mouth o f the
ne st S he worked earnestly a s i f the work was n ow full
o f meaning to her
Disturbed by a pass ing boy S he ew a round came back
and exam i ned he r p rey and returned to mo re d i gg i ng
brought the hopper a foot neare r and propped the th ing up
in a tu ft o f grass and went back d iligently to digging lug
ging out i n he r j aws load s o f earth as large as her head S he
held the masses o f so ft dirt up against the mandibles w ith
he r f ront legs as S he carried i t out backwards to keep the
load from falling to pieces so it looked as i f she were carry
ing a double arm ful up to he r chin AS the burrow grew
deeper s he backed away f rom i t further to perhaps a dis
tance o f two i nches or mo re be fore dropping her load and
occasi onally paused to sweep the whole pi le back with her
forelegs
We examined he r grasshopper a D is s os tei ra c a rolin a
Linn and were aston ished to nd that its hind legs were
gone Evidently they had been care fully bitten o ff though
p robably not by a tr a t u m fo r the cut w a s smooth and not
lacerated as the wounds would have been i f the legs had been
torn o ff
The same day w e found another hopper in the eld w i th
the j umping legs neatly cut o ff B oth o f these mutilated
hoppers displayed all normal activities excep t j ump ing
A fter about ten m inutes more o f digging this one brought
,
62
he r prey neare r to w ith in one foot o f the hole She carried

it by straddl ing i t right s ide up grasp i ng i ts antennae i n her
mandibles clasp i ng her forelegs around i ts neck and strug
gl ing ove r the ground w ith he r two h ind pa i rs o f l egs
The wonder ful m od i cat i on o f the legs o f th i s species as
S hown in gure 36 S hows how capabl e they are o f scrap ing
and carrying the soil as wel l as long enough grace fully to
straddl e and r ide home a huge grasshoppe r A fte r another
brie f per i od o f digg i ng the wasp moved the hopper to the
very b rink o f the hol e an d swung i t around so i t w a s directly
facing the slop ing burrow although i t had formerly been
behind the hole
S he went in turned a round and re
appeared a t once head up seized the grasshopper s antennae
and dragged i t in out o f sight The hol e w a s a clos e t
for the p rey ; i t was w ith di fculty that s he squeezed i t i n
For only one brie f m i nute s he remained inside the chamber
w i th her booty ; then s he scrambled out and scooped i n a few
arm ful s o f di rt arranging each load S l i ghtly but packing
i t l i ttle As the channel lled S he t amped th e d irt down
more S he scoope d i n each a rm ful o f earth w ith a c i rcular
stroke cutting a crescen t S hap ed area out o f th e tiny hill
smoothly and systemat i cally l i ke a workman care fully scoop
ing sand away from one s i de o f a heap ( g
S uch
p rec i sion rathe r startl ed u s since w e were accustomed to
B emb ix w a y o f scratching ove r the whol e region
W ith her head S he packed i t constantly tighter as s he
nea red the top When the depress ion w a s nearly full s he
but b it up chunks o f rm
s to p p ed t a k i n g the l oo se earth
fr esh d i rt wh i ch s he carried i n her mandibles and broke
to p ieces on the S ite j ust as S p hex does w i th l ittle clods
rubbing and p ress i ng down the s o i l w i th them until they
were ground p ieces and became a part o f the rm pack
o f pulveri z ed earth
Jus t as S he appe ared at the mouth o f the hole a fter hav i ng
.
64
one un fam il ia r w i th wasps ways woul d have said at a glance

that her i nterest i n the spot was forever at an end
FI G 38
w hi l
P ri o n o n y x
an c e
p o s i ti o n
p o un d in
of
le
a tr a tum
gd w
o
N a tura l
th e
si e
d e f tly b ra c in g her s el f
s o il
i n her b ur ro
to
w w i th h
h o l d he r b a l
h ead
er
N o te
Then we went back and dug up th e rst h o le for which

S he had hunted so l ong and earnestly expect i ng to nd
what s he S O ca re fully seal ed up i n i t It was stark empty !
Why had S h e been s o very part i cul ar to put the d i rt back
?
i nto al l o f these empty holes
Let us add here that thi s pers istent hab i t o f clos i ng up
the abandoned holes i s not constant We have one record
o f an a tr at u m wh i ch com menced S ix holes be fore s he w a s
sat i s ed w i th a locat i on but made no attempt to k i ck back
the ea rth
The last nest to wh i ch ou r wasp had g i ven such care ful
attention conta i ned the hoppe r w ith the wasp s egg c c
m en ted i n i ts usual position on the hind leg and curved
around the s ide o f the body B ut the D ipterous paras i te
menti oned above must have been too quick fo r the mother
wasp a fte r all fo r fteen m inutes a fter the egg w a s laid
i t and the host w ere al ready teem ing w ith tiny D ipterous
larvae There i s a po ss ibil ity o f course that the hoppe r
may have been parasit i z ed as i t lay i n h i d i ng but thi s chance
,
65
i s sl ight because usually the adult paras ites stolidly igno re

the p rey o f wasp s whil e i t i s le ft lying idl e but take an
interest i n it when they see w ith surety that it i s go ing
to be buried properly
The next day at
p m the hop p e r could still m o ve
i ts mouth parts and legs and when stimulated i t could ut
ter its wings In the inte rval the paras ites had dev o ured the
egg and w ere ente ring and feedi ng upon the body o f the
grassh opper at the point o f attachment o f the wasp s egg at
the base o f the le ft hind leg
We have never found mo re than one hoppe r stored in
the burrow o f P a tra tum W i ll iams found the i r prey in
Kansas to b e a m ature Aul o c ara M ermi ri a neom exi c a na
M ela n op lus di er en ti al i s 52 o r M ta hin a s
On one occasion w e we re so fortunate as to catch the
P a tr a tum i n the act o f selecting he r nesting place
O n that
morning August 30 at 1 0 o clock w e found her s earching
to and fro in the region o f he r grasshopper wh i ch lay a l
ready paralyzed on the gras s We S hould much l ike to
know whether S he had brought i t here from some distance
o r whether s he had killed i t on this sp o t We m ight add
here that the P eckhams are evidently m i staken i n s urm i s
ing that the species rst prepares the nest and then catches
the wherewithal to ll it In eve ry cas e when w e saw the
wasp at work digging w e found the paralyz ed hoppe r near
by This wasp d id not come d i rectly to he r p rey nor did
she at any time wander further than four feet fro m it
Judging f rom he r manner o f hunting here and there over
the region w e thought s he w a s seeking her lost quarry until
a fter exam i ning one bare spot closely for ve minutes s he
began to dig Thi s spot w a s only about eighteen inches
from her hoppe r but i n selecting the s ite s he d id not return
di rectly to her p rey at any time
,
K a n s U ni v
.
Sci
B ul l
: 2 30
1 9 1 13
66
The earth w a s moist and s he dug rapidly carrying out

la rge mouth ful s o f di rt as s he emerged backwards from the
hole dropp ing i t near th e brink and inging i t back unde r
her body and out of the w a y w ith her forelegs When the
di rt had accumulated near the hol e so as to form a S l ig ht
obstruction s he would come out and kick the whole pile
away to a di stance where i t woul d cause no furthe r annoy
ance During the next period we timed he r ; she usu ally
took out ve lo ads o f earth a m inute and oc cas ionally S ix
A fter ten m inutes o f he r fa ith ful digging the hol e w a s
deep enough to i nclude he r whole body ; then her m ind
turned back to he r hopp er and she started out i n quest o f it
H er man ner o f locating i t w a s interesting to watch ; rst
s he walked i n the di rection o f the p rey but a l i ttle to one
side and m issed it ; w hen some d istance beyond i t s he turned
and came b ack but passed i t again Then s he t o ok a S hort
i ght but went too far and la nd ed beyond i t again ; then
another return ight brought her only sl ightly past it once
more and f rom there S he walked a few steps to it S he
mounted it and trui i dled it hal f way to her hole where she
le ft it on top o f a th ick clump o f grass and resumed her
digging for another quarter o f an hour Once more S he
moved he r hopper nearer to the hole ; this time S he had no
t rouble at all in n d i n g i t but walked stra i ght t o it
When
s he brought it near to the hole
w e f elt su re that S he had
the burrow almost ready fo r it for these wasps have a w a y
o f gradually moving thei r booty neare r and nearer as the
wo rk on the burrow p rogresses until i t a rr i ves at i ts destin
ation j ust as the hol e i s completed Why do all o f the
P ri on on yx p ersist i n digging awhile and then bri nging their
prey closer several times be fore its nal burial when they
might as wel l bring it all the way at o n e trip ? D o the
?
wasps grow anxious about the wel fare o f the i r p rey
H ardly or they would bring it at one trip clear to their place
,
68
cinders thrown out on top o f this smooth whit e sur face

I t w as clearly the work o f a P ri on onyx in all p robabil ity
P a tr a tum s ince the only other member o f that genus found
here i s P th o m a e and non e o f them had yet been seen in
thi s V icini ty The gal lery o f this burrow w a s ab o ut an inch
and a hal f long I t w a s tightly packed near the mouth w ith
the mate rial which had be en exc avated I n fo rm and slope
the burrow con formed i n every way to the standard s p ec i c a
tions o f P ri on onyx ; hence w e may conclude that s he nds
the same scheme adequate whether wo rking i n sand or rm
earth The adul t hop p e r i n the cell a D is s os tei ra c aroli na
w a s active in all i ts m embers k i cking vigorously w i th all o f
its legs moving its mouth antennae and abdo men and occa
Dur i ng all o f th i s activ i ty the
s i o n ally uttering its w ings
egg which was glued at the base o f the r i ght hind leg was
not d islodged although it was badly shaken and occasionally
rece ived a full bl o w f rom the middl e leg H ence i t w a s
not i n the least su rpris ing that the egg never hatched and
that th ree days late r it w a s sh r ivelled Th e hoppe r w a s at
that time fa i rly active moving th e hin d l egs mouth and
antennae ; only the rst and s econd p a irs o f legs seemed
helpless
The S ite of the nest was particularly consp icuous because
o f the contrasting colors o f the mound and the adj acent
soil w hi le the tiny black cinders la i d on top o f the gray
clay made the contrast only the more striking In covering
and nishing o ff her burr o w in this way the P ri on o nyx had
merely f ol lowed her usual custom fo r e ffecting its conceal
ment but S he had failed ludicrously by p icking up mate rial
o f sharply contrasting colors
S urely i f th is behavior
was not bl ind i nstinct at l east it w a s colo rbl ind
I n 1 9 1 4 while walking along a country road i n Kansas
one August morn ing w e noticed a hol e in the s i de o f a
wago n rut In a moment the p rop rietor a P a tratum re
.
69
turned and claimed it About twelve inches away and fac

ing the hol e w a s a grasshoppe r a M el an op lus di eren ti alis
Thom 9 [ A N Caudell ] lyin g in the very natural p osi
ti o n o f one at rest When it remained there w ithout mov
ing w e decided that it too w a s a participant in the drama
about to take place
For ftee n m inutes the wasp enlarged the h o le all the
time working out th e dirt w ith her mouth and f o relegs
The mandibles brought o ut the bi g clods whi le the legs
rap idly worked the dust out behind he r Once she paused
to examine the hopper and when S he fo und i t resting per
f ec tly quietly except for the pulsating o f the abdomen s he
at once resumed he r wo rk Fo r e ight m inute s m o re S he
worked at the excavating be for e s he again examined her
p rey S he had mo re di fculty i n nding it thi s time and
walked around in an i rregular ci rcl e twice and a part o f a
third ci rcuit be fore she stumbled u p on i t S he straddled
it as i t lay grasped its antennae i n her j aws stretched her
legs ove r the s ides o f its body and thus trundled it along
j
ust
as
i
n
g
At
a
p
int
about
four
inche
s
di
stan
t
o
(
f rom the nest S he l e ft the grasshopper went to the mouth
o f the burrow poked her head i n turned back and once
more scrutini zed the prey and immediately too k out two
loads o f d irt one from each S ide o f the entrance there
by enlarging the open ing I t w a s di fcult for an observe r
watching th is exact work to th row as ide the conviction that
actual j udgment w as a guiding factor in such l ittl e details
o f behavio r as thi s Then s he continued deepen ing the cav
ity A buzz sim ilar to the shrill buzz o f a y ensnared w a s
occasionally audibl e ; it seemed that each buzz w a s s imul
ta n eo us w ith the energetic p ressing of her head as s he strove
to lo o sen a pellet o f earth At 9 : 1 5 S he started for her
hopper mounted and grasped it the same as be fo re except
ing that this time S he clasped i ts tho rax wit h her f ront legs
.
WA S P S TUDI E S A F I E LD
7O
and struggled fo rward on he r fou r h ind feet Thus they

p rogressed to the burrow There s he laid it down w ith its
head at the very bri nk o f the o p en ing ; but s he le ft i t for
only a moment wh ile she stepped inside turned around and
poked her head out j ust far enough to grasp the hopper by
one antenna and dragged i t i n A fter a br i e f time pe r
haps one hal f m inute she cam e out and immediately com
m en c ed to l l the hole pausing only t o chase a D ipterous
parasite that hove red about wh il e S he was lling i t in H ow
strange i t i s that only the ll ing o f the tunnel seems to stim
ula t e the y s des i re to enter ; S he had had abundant o p p o r
tun i ty to ente r ere this o r even to deposit he r eggs on the
grasshopper a s i t lay wholly unp r o tected but S he seemed to
take no i nterest i n the hoppe r until S he w a s sure th e wasp
had ni shed w ith i t
Wh il e ll ing and pack ing i n the earth the wasp stood on
he r head and p ressed down the s o il p roducing w ith each
stroke the shrill buzz Next s he brought large pellets f rom
a distance and dropped them on the depress ion Then i n
scratching the di rt back s he worked gradually s i dewise w ith
he r face away from the hol e so her path fo rme d a pe r fect
arc o f a ci rcl e the center o f whi ch w a s the nest O ften she
woul d have to go back to the or i ginal spot to get her bear
ings and est i mate exactly where to d i rect her k icks While
thus occup i ed w ithout cerem o ny or w a rning s he aro se and
ew away i n a straight l ine I t w a s th en 9 :30 o r forty ve
m inutes a fte r the time w e d iscovered her
We opened the nest Th e lengt h o f the tunnel d o wn to
the grasshopp er was ve inches The chambe r w a s no w ider
than the rest o f the burrow and lay horizontally No earth
was in thi s chamber and the hoppe r w ith i ts head to the
wall had ampl e a i r S pace around it We dug i t out and took
i t home fo r furthe r observation
I t was still al ive but
seemed severely stunned and for the remainder o f that
.
W A S P S TUDI E S AF I ELD
2
7
Ad am s nds the Tachinid y M etop i a leuc o c ep hala Rossi

hovering Over the i r burrows
T o s ee a P ri on ony x intent upon the herculea n task o f
m aking he r burrow and burying the grasshoppe r one can
hardly i magi n e he r bl ithely re fresh ing hersel f among the
blossoms These wasps se em to feed upon a var i ety o f
plants ; w e have seen them ying f rom ower to owe r sip
p ing the nectar o f blue ve ron ica i ron weed and M el ilotus
and a l ittl e later i n the s ea son early S eptember w e s aw
them w ith extended tongue laz ily Sipping necta r f rom the
orets o f white snake root
O n several o ccasions we followed the clue o f a p il e o f
character i stic chips on the bare ground B y scoop ing away
the sur face so il unde r these mounds to the depth o f an i nch
o r tw o through the plug o f ea rth wh ich the m othe r wasp
had packed rmly i n the mo uth o f the burrow we d i s c o v
ered the nest o f P ri on ony x
The form o f the burrow i s generally uni form sim ilar to
those shown i n the diagram ( g
The holes about
three e i ghths inch i n diamete r go d ownward at a gentle
S lop e and gradually curve until they end i n a horizontal
chambe r
These chambers are not always distinct ; f re
quently they are l ittl e more than a continuati o n o f the gal
lery The burrows are three and one hal f to four i nches in
total length and at the deepest po i nt about one and one hal f
inches bel ow the sur face o f the ground We have neve r
chanced to nd a P ri on onyx bur row o f the fo rm described
by W ill ia m s f a di stinctly L shaped nest about two inches

deep and two and a hal f i nches long There has been ve ry
l ittl e variat i on i n the form o f the fteen nests wh ich we have
excavated
The P ri o n on yx nests which have c o me under our observa
2
'
B ull 1 11 S t Lab N a t H i s t 1 1 : 1 9 5
K a n s a s U n iv S c i B ul l 8 : P I 33 g
.
915
H UNTERS O F LAR G E O RTH O P TERA
73
tion have always con ta ined one o f three species o f gras s

hoppers : D is s os tei ra c a ro li na Linn nymphs Arp hi a
c o ri n a ta S cud
2 large nymphs or M el a n op lus femur
'
rubrum
The rst named w a s the most abundant Riley P ackard
and Thomas record them as pursuing and captur ing the
nymphs 0f the Ro cky Mountain l ocust C ol op ten us s p retus
and Adams ( loc c i t p 1 9 5 ) al so says they use M ela n op lus
femur rubru m The p rey was always placed in the oval
terminal chambe r w ith its head toward the wall away fro m
the exit a w i se precaution which would p revent the hop

e
r
s
escap
in
cas
e
the
stinging
S
hould
be
imper
fectly
done
e
p
The upper portion o f the channel was always lled w ith
earth and nea r the sur face this was packed so rmly as to
be indi stingui shabl e from the surround ing so il The hori
z o n t al chamber at the end w a s not lled w ith earth but con
ta in ed only the grasshoppe r surrounded by plenty o f a i r
space May i t be that thi s terminal chamber i s made i n th i s
form so t hat the di rt wil l not roll in and pack around the
grasshopper ?
The faith fulness o f these wasps i n returning to the un
ni shed nests i s variable We have described above the case
o f one wasp which spent a whol e hour i n dil igent search for
an unn ished hole only to l l i t up and desert it the m o ment
S he found it
On one other occas ion w e disturbed a wasp as
she w a s making her nest and found the next day that s he
had returned and n i shed i t In another instance w e met
exactly the op p o site behavio r : the wasp w ith he r prey a
large D is s os tei ra c a rolina right bes i de her w a s putting the
nish i ng touches to her burrow when our foo tsteps fright
ened her away S he returned and found the hol e agai n b e
fore we passed o n but when w e came back that a fternoon
.
4
5
d ly i d en ti ed b y M r A N
U 8 B ut Com m 1 8 7 7 : 31 8
All
.
ki
.
C a ud el l
74
to dig up the n i shed p roduct w e were sur pr i sed to nd

that S he had deserted i t j ust as w e had l e ft it that morning
The nest remained sl ightly un n i shed and the hop p e r nearby
was be ing di smembe red by ants
Two other unnished and deserted burrows o f P ri on on yx
probably a tra tum were found On e w a s ev idently ready for
the closing ; even the hopp er a M el an op lus femu r rubrum
w a s i n place at the bottom but the owner w a s gone the d i rt
w a s d ry and the l o cust was be i ng carried aw ay i n b its by
ants The o ther w a s almos t as fa r advanced ; the p rey an
Arp hi a c ori n a ta 9 lay di rectly i n front o f the hol e and fac
i ng it the l ast position be fo re the wasp drags i t i n but

th i s too w a s being plundered by ants There i s o f course
a poss ibil ity that both o f these failures had been caused not
by the fa ithlessness o f the moth er wasp but by some tragedy
wh i ch had overtaken he r We m i ght ad d here that the
mothers themselve s a re not exempt f rom enem i es fo r P i erce
reco rds thi s species as be i ng stylopi zed by Op htha lm o c hlus
dury i and i n o n e instance where a mal e paras ite emerged
three days a fter the death o f the host
On o n e Augus t morn i ng we w ere interested i n watchi ng
the courtsh i p o f a P a tra tum At 8 :4 2 a black fem ale was
seen at rest on a small M eli l o tus A fter a few seconds a
s econd i nsect appeared ew d i rectly to the rst mounted
and t r i ed hard to e ffect a un ion and ew o ff During the
next fteen minutes thi s per fo rmance was repeated about a
do zen times ; occas ionally the male would y to a distance
o f fteen feet re st fo r a second and then return t o the female
and resume act i v i ties but more o ften the ights were S ho rt
The male woul d hover to and fro in a space o f about three
inches i n the ai r in f ront o f the femal e be fore mounting ;
then he would curl his abdomen beneath hers Mating prob
ably occurred dur i ng thi s coquettish per formance but i f s o
,
B ull U
.
N a t M us N o 66
.
38
1 909
6
7
as w e watched her S he deserved our high est respect for

faith fulness to duty
Her method o f di gging was s imilar to that o f P a tr a tum ;
s he wou ld plunge into the hol e head rst grab a lum p o f dir t
or a c i nde r b a ck out and drop her load a few inches away
When the bo re was about hal f completed s he found he r
hopper w i th l ittle di fculty mounted it again grasped its
antennae in her mandibl es as usual and away S he w ent ( g
S he d id not exactly y but she made such rap i d
speed that s he must have ai ded he r progress by beating her
w ings S he le ft the locust about twelve inches from the
hol e unti l s he enlarged it Then S he returned again to h er
p rey and desp ite the fact that S he w a s very near to it she
found i t w ith more di fculty than o n th e prev i o us occas ion
I t took he r one m inute to d iscove r i t and at one t ime s he
passed it when only one i nch f rom where i t lay I n the
usual manne r s he mounted it grasped the hopper s antennae
i n he r mouth and d ragged i t to w ithin th ree inches o f the
burrow H e re s he evidently decided that it was a trie larger
than the hol e wo uld yet adm it so s he l aid i t dow n and t e
moved thre e mo re mouth ful s o f di rt S he then carr i ed the
p rey to w i thin one i nch w ent i nto the hol e once more to
make sure that all w a s ready came out again and dumped it
head rs t hal f w a y into the op en ing ; then S he hersel f
crowded i n ahead o f i t and dragged i t i n a fte r her S he
remained i n only o n e hal f m inute came out and began to
ll in the di rt A fter this f o r many m inutes she ca rried
large cinders and placed them ove r the s ite The heavy cin
de rs were moved w ith the mandibles w i th comparat i v e eas e
S ixteen m i nutes later s he stopped p i l i ng cinders and for
two m inute s walked an d too k S hort low i ghts about
the nest survey i ng the s ituation ; then s he came back to the
,
T hro ugh
g ro un d
7
an erro r
i n the
gu
re ,
t he m i ddl e l eg
f ai ls
to
r eac h
th e
H UNTERS O F LAR G E O RTH O P TERA
77
very spot and critically added more cinders took a few more
last looks and nally at 5 2 9 she ew o ff and rest ed on the
grass some distance away as i f all he r interests were now
n d the past w a s a closed book
It had taken
i n the future a
,
FI G
40
Hom e
to the b u rro
d b Oun d P ri o n ony x thomae
ar
E x ac t
si e
tra n s p o rti n g h e r l oc us t
her j ust f ty four m i nutes to di g he r burrow bury her prey

lay the egg and conceal the s p ot
A fte r s he had gone w e opened the nest and found the
hopper w i th its head to the wall The chambe r w a s only
one inch underground and one inch long o f no den ite S hape
but it was as wel l constructed as i t could be in such rough
material We found the brown egg on the right femur nea r
where it j oins the thorax We took the egg and its host
home and guarded them care fully The next morning July
1 5
at 7 o clock w e found the legs antennae and mouth
parts o f the hopper active and excrement had been passed but
the femu r upon which the egg re sted was immovable With
what del icate accu racy the l ittle mothe r wasp had done he r
work ! B y the following day the egg was slightly enlarged
and had changed from brown to green The morning o f the
1 7 th found a larva fat and large still clinging to the femur
o f the hopper and eating away the thorax near the j o int
-
8
7
July 9 revealed a light brown coc o on j ust c ompleted ; the

hopper had all been eaten O n August 1 5 scarcely one
month later a p er f ect adult f emale emerged Th is S hows
that in this species there i s more than one generation each
year
H artman in an inte resting account o f thi s species tells
o f how he accidentally ran into eight nests al l i n a space not
larger than hal f o f th is page

Th e chambers were so close
togethe r that some had but a quarter o f an inch o f wall b e
t w een them H e says the chambers wer e oblong about t w o
inches in length and one hal f to seven eighths inch across and
two inches below the su r face o f the earth
Wi ll iams observed two specimens that secur ed locusts
belonging to the gene ra Am p hi torn us and Aul oc a ra
Another chapter in the l i fe o f P th om a e the hunting o f
the p rovis ions must b e taken f rom another individual
A terric commotion i n the l ow gras s attracted our a t
tention An adult g ro s s hop p e r Arp hi a x a n thop tera B urm
A
c
N
Caudell
lay
on
its
ba
k
v
i
olently
apping
its
w
ings
]
[
against the earth as i f in great d istress A l ittle red lump
on the ventral S ide o f the tho rax expla ined the mystery ; it
was the abdomen o f a P tho ma e w ith its sting buried deep
in the hopp er s thorax ; the rest o f the body o f the wasp was
curved around the grasshopper s S ide The struggle was
more violent than that o f an insect on its back trying to
right itsel f ; besides a g rasshopper seldom has much d if
culty i n turning over i f it happens to become invertedi n
fact they are seldom s o cl umsy as to get i nto that p redica
ment Th e po int is thi s acti on w a s s imply an aimless and
con clusive muscular contraction a w rithing resulti ng from
the sting o f its enemy A grasshopper i s s o seldom seen i n
thi s position that the wasp must have e it her attacked i t on
1
B ul l U n iv T ex N o 65 : 64 1 905
K a n s U n iv S c i B ul l 8 : 22 7 1 9 1 3
.
80
A fte r wa i ting i n va i n for he r return w e l ooked at the

hopp er again ; i t had somehow r i ght ed i tsel f and sat o n the
ground with i ts w ings closed Closer exam i nat i on showed
that it w a s powerless to use its legs but it mov ed its w ings
i n resp o nse to stimulation I t tried to walk but the only
result o f i ts e ff o rts w a s a sl ight quive r o f the l e ft h ind leg
We p i cked i t up a l i ttle later to exam i ne i t more closely on
the open hand Imag i ne our surp ris e when w ith the rst
pu ff o f w i nd i t spread its w i ngs and ew a di stance o f
forty feet and w a s lost i n the v egetation E v iden tly t he legs
had b een properly paralyzed but the w ing ne rves and
muscles w ere una ff ected Th i s c o ndit i on i s usually s uf
cient by reason o f the fact that hoppers cannot normally use
the i r w ings fo r ight w i thout rst l eap i ng i nto the a i r by
means o f the i r ump ing legs A s prey they m ay occasional ly
escape by som e unusual ci rcumstance as i n th i s cas e but
such cases a re so rare that the re i s n o need o f the wasp bur
den ing h e rsel f with e ither the i nstinct o r the task o f p a ra
ly z in g the W i ngs o f her p rey
As to th is hop p er it p robably
l ive d and grew fat even i f i t w a s unable to use i ts hind
legs but w e should l ike much to kno w whether the inj ured
members fell o ff We have seen healthy hoppers w ith the
hind legs s o neatly trimmed o ff at the ba sal j oint that w e
wonde red how they came so ; may i t be that P thom a e o r
P a tra tum had once had them ?
,
The Relat i o n o f
S ti z us
P mon ony x tho m a e
un i ci n c tm
Say [ 8 A Rohwe r ] to
.
F ahr
While Chl ori on c y a n eum w a s working on the c i nde r bed

as later described a P ri on onyx thom a e appe ared r econ
S he at once w ent to work ener
n o i t e ri n g i n the same el d
i
ll
and
dug
a
hole
a
hal
f
i
nch
deep
then
deserted
i
t
a
e
t
c
g
y
-
8I
and attempted another at a spot ten feet distant and aban

domed that also ; a l ittl e later s he commenced a th i rd burrow
about ve feet f rom the second This last location seemed
to o ffer more favorable conditions He re the wasp worked
dil igently among the cinders and in ten minutes had dug
a hol e the l ength o f he r body w i th an open ing one hal f
inch w ide This w a s good w o rk cons idering the di f culties
o f d igging i n packed cinders It seems that this spec i es has
a l iking fo r cinders for the other that w e saw worked in
the same material desp ite the fact that only a small p ropor
tion o f the available area w a s covered w i th it
The blue wasp C cy an eum wh i ch wa s forag i ng under a
brick p ile n ear by annoyed her twice by entering he r open
burrow ; but s ince s he entered every hole and crev i ce i n the
brick pil e and cinde rs thi s w a s p robably o nly an accident
This P thomae foll o wed the same tec hn i que i n he r work
as P a tra tum :s he w ould carry up the so i l c i nders etc w i th
he r mouth back out w ith the load and drop i t near the or i ce
w ithout turning around and run i n again hea d rst so o n
to emerge i n the sam e mechanical manner l ike a l ittl e toy
wound up w ith a spr i ng When the deb ris had accumulated
so as to be annoying she p a used long enough to rap i dly kick
and b rush it all together i n a n eat l ittle heap some di stance
away
A fter s he had dug down about the distance o f he r own
body length she turned to a h ori zontal di rect i on and ex
cavated the chamber Thus the ceil ing o f the chambe r
when compl eted w a s only about a hal f inch beneath the
sur face o f the ground The open i ng was la rge enough to
pe rm it us easily to s ee her moving about inside
A fter a hal f hour s arduous work she came out o f her
b urrow and slowly walked about in the i mmediate vicin ity
o f her nest We thought s he was getting he r landmarks
p reparatory to br i ng i ng her booty home H er sl ow and
.
82
del iberate actions indicated thi s but presently she ch anged

her m ind and again went into the hol e to make certain i m
m
r
n
o
v
e
e
t
s
in
vi
ew
o
f
the
com
ing
storage
remov
ed
a
f
ew
p
mo re mouth ful s o f dirt and again strolled slowly about her
nest Then s he ew o ff for he r hopp er D uring th is period
o f he r wo rk w e had made a di ligent search fo r her prey
which w e we re certain s he had in h iding somewhere near
B ut now w ithout hes itation or uncerta inty s h e walked di
rec tl
to
a
brown
br
i
ck
on
the
ground
at
the
s
ide
o
f
wh
ich
y
lay he r hoppe r almo st i ndistingui shable agai nst its similar
colored background
Once while s he was a t work in the hole a red banded
S ti z us un i c i n c tus had po k ed he r head in and walked away
w ithout further intrus ion ; but now while s he w a s gone the
same meddler return ed and wandered about the vicinity
w ith a loo king for something manner

P tho m a e straddled the hopper graspe d its antennae in
he r mouth and struggl ed on he r way ove r the roll ing cin
de rs I t was w ith conside rabl e di fcul ty that s he rod e the
hopper ; bes ides the grasp on its antennae s h e gripped the
hopper s neck w ith he r fo relegs ; so she walk ed w ith her
hind legs wh ile the m iddl e l egs attempted to ass ist but
owi ng to th eir shortnes s o r the thickness o f the burden they
seldom reached the ground The hoppe r i tsel f o ften ass isted
in their progress i n th i s way : as tho m ae d ragged he r prey
forward the tars i o f its h ind legs woul d cl ing to the rough
cinders and whil e the heavy body wa s be ing pulled fo rwa rd
the hind legs woul d thus be stretched out to thei r fullest
extent ; o f course th is process w ould l i ft the rear part o f the
grasshopper and make the body pitch fo rward When the
legs had reach ed the i r full extent they would snap shut and
the process woul d be rep eated push i ng the heavy b ody
sl i ghtly forward each time thus at least avoiding h in
,
84
to remove p roj ecting into the galle ry ; this o w obstructed

the way s o s he had troubl e i n getting her pr operty i n but
s he worked strenuously ins id e the hol e and w ith our assi st
ance on the outsi de worked her locust i n a fte r ve m in
ute s labo r Agai n S ti zus w a s seen lurking about i n the
vicin ity o f the n est
P Thoma e came out and lled her hole care fully rst by
carrying in mouth fuls o f so il and precisely arranging i t on
the inside and then kicking mo re di rt i n and going i n a fter
i t to pound i t down mak i ng a loud hum as she p ressed w ith
all he r strength against the tighten ing so il I n getting di rt
to ll i n s he rs t to o k that nearest the hol e and when thi s
was gone instead o f taking all o f the earth wh ich s he had
kicked into a h ill s he dug at the foo t o f thi s tiny h ill and
hollowed o ut a sem i c i rcular space j ust a s w e hav e seen
P a tra tum do ( g
O f cou rse a s s he dug at the foot
o f the p ile s he unde rm i n ed i t and much o f th e d irt tumbled
down and rolled i nto o r near to the cav ity w here s he needed
i t When the hol e w a s hal f covered she w a s aga in v i s it ed
by S tizus but she at onc e gave chase and th e meddl esome
intruder ed The task o f ll ing and cove ring w as con
t i n ued an d wh en th i s w a s done several cinde rs large in
s iz e but o f l ight weight ( see g 4 2 exact siz e ) were
b rought i n th e mandibl es and placed ove r the spot A col
lection o f twelve such cinders was taken f rom her mouth
with the fo rcep s as s he b rought them to the place ; th is
shows how tame s he w a s or how accustomed to our presence
s he had grown
She surrendered th e cinde rs to us peace
ably and w i thout deance ; howeve r w e did not pull them
away f rom her but merely held th em rmly w ith th e forceps
s o s he could not move them further So she gave them up or
she m ay have merely thought that they we re too heavy or
A fte r a f ew mo re k i cks
to o tightly lodged for her to move
s he abruptly ew away
,
'
H U NTERS
FI G
of
A
42
th o m a e
.
c o l l ect i o n
an d
OF
LAR G E O RTH O P TERA
o f c i n d ers ,
d ep o s i t e d
on
x a c t s iz e
t he to p
of
c a rr i e
85
d i n th e m a n dib l es
he r b ur ro
Our esh and bones were weary o f sitting on all fours

on the cinders for two hours and i t was grow ing late but
sti ll w e we re eager to see i f S ti zus would show any furthe r
i nterest i n the nest We waited fo r ve m inutes and she did
not reappear so w e began to dig up the nest and had re
moved enough so il to expose one o f the hopper s tars i when
behold ! S tiz us appear ed upon the scene We sl ipped back
and s he came neare r and nearer to the nest by walking z ig
zag fol low ing almost arcs o f circles around the hol e w ith
her head close to the sur face and antennae vibrating until
S he located i t apparently w ith less
s he came to the nest
di fculty than wasps sometimes expe r i ence in locating the i r
She approached i n the manne r o f one fam iliar
o w n nests
w ith the s i tuation and hav i ng bus iness there She k i cked
away much o f the ll ing until s he had remov ed the hard
plug and arr i ved at the l oo se mater i al beneath ; then she
,
86
wo rked he r way into the nest at the right s ide 5 f the hopper
remained in for about ve m inutes and eme rged at the le ft
side show ing that at least s he could turn around ins ide the
chamber S he went in aga in and thi s time w e removed a
l ittl e o f the soi l yet coul d not s ee fully what s he w a s doing ;
but thi s much w a s cl ear that she was standing on the hop
pe r w ith he r head ne a r th e fore part o f th e insect quietly
s itting there w ith the abdomen pulsating for over v e m in
utes S he came out and kicked i n all o f th e loose dirt then
dug up more w ith he r mandibles k icked it unde r he r body
into place and p icked up and placed a few more bits until the
hole w a s once more nicely c o ve red S he then went a few
feet away and calmly sat down to rest when s he was taken
to secure he r i dentity
We dug up the ne st and found a hole o f the typ e usually
made by both P a tra tum and thom a e only the chamber w a s
unusually long in th is cas e The hoppe r lay ther e w ith its
head to the wall The egg o f thom ae was i n its usual pos i
t i on on the r i ght femu r at th e base o f the abdomen but 10 !
it was only an empty sac ! Evi dently S ti z us had mutilated
the egg by malaxation or had sucked the contents fo r food
We found no S tiz us egg ; but i t i s pr obable that i n the d ii
culty of opening the nest in a harsh cinder bed i t w a s
knocked off by crumbl ing walls We know that S ti zus di d
not de p o s i t an egg during he r second stay in the chamber
for w e could s ee he r abdomen all the while but we suspect
that s he w a s then mutilating the egg o f thom a e a fte r prob
ably hav ing lai d he r own
S
We wonde r by what sens e ti z us found the nest To be
sure s he had returned to the spot while P tho mae w a s at
work su fciently o ften to get the landmarks o f the v icinity ;
but consideri ng the remarkable way i n whic h the nests are
cove red and conceal ed w e w onder how she c o uld nd the
,
8S
ove r the grassy part o f the eld exam in i ng b are or rough

ened spots j ust l ike the one described above S he m o ved
calmly and w ithout haste or agitation A fter examining
about ten sp ots in a coupl e o f m inutes s he a rrived at a
certai n inconsp icuous p ile o f loose earth in outward appear
ance very s imilar to all the others Instantly her manner
changed S he became greatly excited ne rvous qu i vering
w i th eagerness S he p robed here a n d there in the p ile unti l
i n a f raction o f a m inute s he had loc ated the l led up hole
beneath one s ide o f the p i le and then eagerly began digging
S he burrowed steadily and rapidly in the f reshly stirred
ea rth and pushed and ung the loos e d i rt out between he r
l egs and under her body S he worked w i th th e fu r i ous
eagerness o f a dog digg i n g o ii t a rabbit ; surely i f s he could
have barked or snorted she would have done so I n a m in
ute or s o s he w a s through the plug o f earth and disappea red
i nto the cell We waited bre athlessly knowing that th i s
w a s the t i me when s he w a s probably crushing the P ri on on yx
egg and lay i ng her own but w e nearly fell on our backs
whe n she came out a t o n c e washed her fac e thoroughly
rubbed her legs vigorously and unceremon iously ew away
leav ing the hole w i de open ! The re w a s no indicat i on o f
he r retu rning or giving any more attent i on to the place
s o w e dug i t up to see i f it really was a P ri o n on x hole and
y
i f she had done anything The case was quickly expla ined :
the hopper i n the P a tra tum chamb er w a s reeking w i th
paras ite larvae but bore neithe r P ri on ony x no r S tiz us egg
u
Wis e S ti z s
Th e D i ptera larvae had p robably devoured
the P ri on onyx egg and the S ti z us knew at a glance that
the same fate would be fall he r babe i f s he l e ft i t there so
she ed fro m the place at once Wher e will adaptations and
interrelations cease !
A f ew m i nutes late r w e n oti c ed s everal paras i t i c D i ptera
'
'
89
hover i ng around the wreck o f the tunnel although both

?
n
hopper and wasp were o w gone H ow did they know
It
d i d not any longe r loo k i n the l east l ike a P a tra tum hole
all dug up a s it was D id som e P ri on on yx odo r r emain on
the di rt which s he had handle d by wh ich the paras ites and
also S tiz us could detect th e location ?
On one oth er occas i on w e opened a carelessly clos ed or
as w e late r concluded a newly o pened burrow which w e
too k to be that o f a P a tra tum but were surprised to nd
beneath the rumpled earth cover i ng a f reshly kill ed hopper

and i n the chambe r w ith it a l ive S ti zus What business s he
had there w e can only surmi se
To be sure the above cases give only ci rcumstantial evi
dence wh i ch i s generally not accepted a s proo f in the courts
o f law Yet w e think that the evidence here i s strong enough
to j ust i fy our i ndict i ng S tiz us as a cow bi rd was p and one
h ighly skilled i n her pro fession In thi s suspicion w e are
not alone f o r Will iams al so ha s suggest ed :
While the evidence at hand i s i ncomplete it seems more

than p robable that the common red banded b em b ec id wasp
S ti zus un i ci n c tus Say pl ays the part o f a burglar and use s
the locust captured by P ri on ony x a tra ta as food fo r he r o w n
young Un i ci n c tus is a rathe r compact insect som ewhat
in ferio r i n s i ze to and les s powe r ful than the sphec i d
I n S tanton county one o f these wasps was observ ed to hover
about a f reshly made tunnel apparently that o f a P ri on o nyx
which i t ente red while the sphecid w a s away Th e latter
had brought an Aul oc am nea r this burrow which be ing
occup i ed by S tiz us wa s nally deserted by the d isgusted
,
P ri on on y x
I n Mo rton county July 7 1 9 1 1 I came upon a S tizus

un i c i n c tus engaged i n smoo th ing o ver a spot with her feet
l
l
d
up burrow which in
e
I unearthed what p roved to be a
,
0
9
form and d imens ions resembled that o f a P ri on ony x I n

the single cell lay a M ela n op lus B ut wh ere the P ri on on yx
egg was to be expected on thi s egg w a s only a small bi t o f
so ft matter p robably the remnant o f a sphecid egg de
stroyed by the S ti zus w hile c ep ha la d o f thes e remains was
a short wasp egg doubtless that o f S ti zus
Certainly the short legged S tiz us un i ci n c tus does not
appear to be a s ui c ien tly powe r ful insect to capture and

subdue l oc usts o f the s iz e and v i gor o f those wh i ch serve a s
the prey o f P ri on on y x
I f we conside r thi s i dea tenabl e i t may throw some l ight
upon the ci rcumstances wh ich obtai ned when once we fo und
in d i ff erent places th ree o p en holes appa rently o f P a tra
tum
We ope ned them further ; one hopper had been
almost al l carried away by ants ano ther was heavily para
and th e th ird
s i t i z ed by D ipte ra larvae and nearly riddl ed
had both ants and parasitic larvae
H ad S tiz us opened
?
these and le ft them open l ike the one described above
We
have at other times al so been mysti ed by open P r i on on yx
holes o r thei r grasshoppers ruined by ants
S om e such
cause must b e behind thi s waste fo r it i s too frequent to
be attributed to mere accident Furthermore when P ri
she doe s i t s o tho roughly that it
o n o n y x clo ses her hole
takes a sneak thi e f to get in
S tiz us undoubtedly o pens the nests for the pu rpose o f
layi ng her o w n egg True we have n ot found her egg
the re even though the P ri on onyx egg had been destroyed
but this may well have been due to our faulty m an ip u
lation i n digging I t seem s hardly poss ible that the taste
o f S tiz zts can be so perverted that s he cann ibal like goes
into the nest on ly for the purpose o f sucking the contents o f
th e egg f or we nd s he i s a nectar f e eeder We have o ften
seen her feeding upon the o wers o f white sn ake ro o t
.
'
'
2
9
time so why did they not come about be fo re the hol e w a s

opened ? P robably j ust because the ies could not scent the
hoppe r through an inch o r mor e o f well p acked earth S o
i f the y w ith its highly special ized of a c to ry organs
cannot scent the contents o f the n est when i t i s cl osed w i th
earth how can w e have the audac i ty to suspe ct that 1 mi
c i n c tm nds he r way to the burrow s by the sense o f smell
?
alon e
O f cours e you may say it is poss ibl e that mother
P ri on on yx wh en carry i ng o ut the earth leaves he r ta i nt
u p on i t P e rhaps s o but even i f she does i n th is c ase the
condition o f th e hoppe r and paras ites show ed that the nest
had b een sealed f or at l east thre e days and surely any
taint that P ri on onyx could le ave at her doo rway would in
three days o f wind and weather have b een reduced to a
m in i mum G ranting that P ri on o nyx does leave an odo r
w e would aga i n have to prove that S ti z us has the p o wer to
perce i ve i t i n s o del icate a degree I t i s hard to bel ieve that
s he locates th e n est by the grasshoppe r odo r
fo r the el d
i s at that time o f yea r constantly overrun by ho rdes o f
grasshoppers and ce rta i nly s he cannot by scen t p ick out
a bur i ed one which i s j ust l ike all the others except that
i t i s paralyzed
Thus i t seems f rom the evi dence at hand that the greater
part o f the respons ibil ity in d i scovering the hidden nests
falls upon th e sens e o f s ight although it i s almost beyond
b el ie f that any on e sens e coul d be d ev elop ed to such a de
gree o f sens it iveness
The highly speciali zed i nstinct o f S ti mts to get the
hoppe r only a fte r it i s bu ried i s w o rthy o f note Th e hop
per l i e s fully exposed for a long p e riod aw a iting bu rial
during w hich tim e S tiz us igno res it but susp i ciously v isits
and revisits the hole Would i t not have been less compl ex
i f her instinct had been s o developed a s to lead her to seek
.
93
the h opper as it lay and depos it her egg and then and the re
get i t out o f her system both phys iological ly and psych o
logically ? B ut no the chance o f inj ury to her egg i n the
transportation o f the hoppe r f or burial and the que stion
o f the survival o f her young o n e in a tussl e w ith the in fant
P r i on on yx p roves he r w ay sa fer a fter all and s he i s r ight
i n wanting to s ee the food sa fely in place the a tra tum egg
destroyed and the host devoid o f paras ites be fore she w ill
take any chance s w ith her own young
,
i c hn eum o n eum
C hl ori on ( P ro teros p hex )
While walking ove r the sparse grass on a clayey plate a u

w e scared up a large sand wasp Chl ori on i c lm eum o n eum
I t created a great commotion buzz ing about our heads in
threatening manner until w e w ithdrew I t then returned
to its bur row wh ich was near by and resumed its digging
The hole w a s al ready nea ring completi o n ; it went straight
down i nto the ground from an o pening one fourth inch i n
diamete r About four inches di stant w a s a large pile o f
which had been taken from
pellets o f earth ( g
the hole
P resently s he stepped aside to get he r prey wh ich she
w a s from time to time bri nging neare r to the hole ; it was
a long horned green grasshoppe r Th en a fte r the usual
manner o f certain wasps she lai d it down and went inside
once more f or the nal survey o f the pocket came out and
brou ght the hoppe r to within one hal f inch o f the open
ing ; s he went in head rst turned around and poked her
head out j ust far enough to gras p the hoppe r s head and
then dragged i t down out o f sight A minute late r s he
appeared at the sur face for j ust a moment and at once went
down and rema ined ins ide the burro w for three m i nutes
,
94
When s he emerged ag a in s he ew aw a y d irectly leaving

the h o l e w ide open Th is w a s at 1 2 30 p m We thought
s he had gon e fo r ano t her hopper ; but when s h e did not
return by 3 o clock w e ope ned the nest The hole went
straight down into the hard so il fo r s even inches then it
tu rned at a right angl e and in the direction j ust opposite
to the p ile o f p ellets continued for three inches and ter
m in a ted in an enlargement or pocket ( see g 4 3 one th ird
natural si ze )
The
p
cket
in
th
is
one
a
somewhat
lar
o
s
w
(
ger than that illustrated by th e P eckhams ) I n it were four
green long horned grasshoppers Orc heli m um mtlg ar e one
o f wh ich bo re a young larva w hich appeared to b e a D ip
Anoth er female nymph wh ich w a s larger
t e ro us parasite
body
length
e
eighths
inch
bore
a
yellow
sl
i
htly
v
g
(
)
cu rved egg th ree s i xteenth s i nch i n l ength The other two
hoppers w e re males measu r i ng nearly three quarte rs o f an
inch i n length They were not active but were still able to
move the antennae and mouth parts when stimulated
B y the evening o f July 2 8 tw o d a ys later the was p la rva
had grown enormously It had eaten the contents o f three
o f the green hoppers and was greedily working ove r the
garbage a few heads and l egs and actually eating the so fter
parts o f the integument so w e gave i t the f ourth g rass
hoppe r S hortly a fter thi s i t spun a we a k and i ncomplete
T w o weeks later w e found that this had been
c o coon
ravaged by that p esti ferous intrude r M eli tto b i a
On the sam e day and in the same place a s econd burrow
Th is had th e same external
w a s di scovered n ea r the rst
appearance a hol e v e e i ghths inch in d iamete r running
straight downward and a large neat p il e o f loos e dirt and
small pellets four inches away The wasp soon retu rn ed
f rom her ight o f alarm and resumed he r digg m g S he would
walk in head rst and c ome out backwards and cont i nue to
,
'
WA S P
6
9
S TUDI E S AFI ELD
i ntently s he seemed agitated and du r i ng the hal f

hour that
we watched he r s he nervously ew away ve times for
no cause that we could perce i v e
We coul d not wa it that day to s ee her nish the wo rk
s o w e retu rn ed to the spot two weeks late r when w e ex c a
v a t ed the n est and found the pupal case
The hole w a s
exactly l ike the one in gure 4 3 g o i n g strai ght dow n for
seven inches then turn ing squarely i n th e direction oppo
s i te the d i rt p ile to form a chambe r
Another nest o f Chl ori on i c hn eum on eum f ound i n the
sunny s ide o f an open shed at Lak e Vi ew Kansas had two
c ells about s even o r e ight inches be low the sur face o f the
ground The po ckets were h ori zontal and sealed a b out an
inch and a hal f l ong b y one i nch w i de B oth cells had bee n
made f rom th e one gallery and both contained long ho rned
hoppers o f the speci es Gremli n mm c a lcam tum R
H [A
N Caudel l ] s ix i n each p o cket The orice and the main
burrow o f the hole w ere on e hal f inch i n diameter The
hoppers ve m ales and one f emale w ere i n o n e cell and th e
egg w a s fastened to one ventro laterally between the rst
and se c ond legs ; i n the other cel l w ere three m ales and three
females and the egg wa s on the ventr a l s ide o f one B arth
nds that th i s was p has several cell s leading o ff at right
angles to the main passage and that i t store s i ts nest w i th
as many as twelve grasshoppers
Fo r thi s species Hancock s notes substantiate the ob s er
In addition he found the prey
v a t i o n s o f the P eckham s
to be females o f Orc helzm um deli c a tum and O vulg are and
both se x es o f Con oc ep halus a tten ua tus H e found one wasp
to weigh 5 grains and fou r hoppers weighed 7 1 0 1 1 and
One wasp m a de t w o nests i n succession close
1 0 grains
together
1 B ul l
W i s c N a t H i s t S o c 6 : I 34 1 9 08
,
11
1 2
'
1 2
N ature S tudi e s i n T em p erate A m eric a
95
-2
01
91 1
97
The p rey from ou r tw o nests survived the ir sting and

imp risonment equally wel l The next day in one cell one
had rotted on e h ad died and f our res p onded to st imula
ti on ; in the ot he r c ell one had rotted on e responded to
stimulation and four w ere motionless but app a r ently living
Four days l ater all w ere dead
These C i c /
m eum on eum als o are annoyed by various
pests
The eve r pres ent M eli tto bi a we re found in thei r
pupal cases kept at h o me although in nature when the un
d erg ro un d nest i s closed it i s imp robable that they can
reach them A hoppe r taken f rom one o f the i r nests had
a larva attached to i t that w a s probably a Hymenopterous
parasite Our attempt to bring i t to maturity fa iled
We have noticed these wasp s feedi ng on owers both
goldenrod and sweet clove r Robertso n reco rds them as
feeding upon the owers o f var i ous s p ecie s o f S oli dag o and
Cockerell nds that i n New Mexico they frequent the
owers o f C l eom e s errula ta Fol som in hi s Entomology
gures an insect o f thi s s p eci es with p o llen ia o f m ilkweed
attached to its l egs
Fernal d nds that they v is it the
owers o f sumach clemati s asclep i as m int etc and says
that the species has p robably the w idest di stribution o f any
o f th e Chlo rioninae i n America H e has s ee n specimens
from various states in the Un ited S tates and gives re fe r
G uatemala N ic a ragua
en c es o f the i r occurrence in B raz il
Costa Ri c a P anam a G ui ana Venezuela Cuba Jamaica and
Santo Domingo H oward say s that thi s species is grad

ua lly extending its northward range
P ierce nd s thi s i nsect to be the host o f the tw isted wing
parasite S trep s ip tera Adam s says that the larva o f th is
species i s probably devoured by the parasitic y M eta/
no
.
'
1 8
1 3
1 4
P ro c
Ci r
U S N at M us 31 : 402 1 907
B urea u E n to m 9 7 : 2 1 908 F o ot
n o te
8
9
Howard gi ves a goo d gure o f thi s wasp i n

Pl 5 , g
.
T he
I nsect
B ook
Dav i s found the follow i ng p rey l n the nests w hich he

opened : Atlarn ti c us d ors alis three in each o f tw o nest s and
ve i n anothe r and elsewhere he sa w a wasp carry i ng a
C on oc ep ha lm tri op s P ackard i n hi s G uide says they use
Our record s mention
Orc hili m um w rlg ar e o r O g raci l e
only Orc heli m um i n thei r nests All o f these are the long
ho rned grasshoppers fo r wh ich they show a p re ference
15
'
'
Locust hunte r
that M akes Tw in Cells
All summer long w e had scoured th is ba re sp ot i n the

eld almost daily ( g
yet i t w a s the m iddle o f S ep tember
be fo re w e d iscovered the rst burrows o f an unknown was p
The mouth o f the hole open i s about th ree s ixteenths

i n ch i n diameter clear cut and m ight eas ily be mi staken
for a beetl e s o r sp ider s ho le wh ile the neat l ittle mound
o f d irt nearby i s almos t i ndi stingu i shabl e f rom the dainty
littl e ant h ills wh ich abound i n the eld S p hex p i c tip en n i s
carr i es out and p il es up the di rt i n the fo rm o f l ittle ch ips ;
P ri on o ny x and B em bix k ick i t out as dust but th is species
br i ngs it out and p i le s i t up i n granular form only a tri e
coarse r than that o f the tiny ant s S he mu st understand
the art o f keep i ng hersel f modestly i nc o nspicuous and must
work w ithout all the bombasti c commot i on i n which B em b ix
indulges for w e hav e n eve r been abl e to catch he r at he r
work
s ed our curios i ty on
One o f these p iles o f g ran ul es ai o u
S ep te mber
We exam i ned al l the earth about i t ca re
fully but could detect no trace nor scar on the gro und w he re
the soil had been d isturb ed s o w ith the trowel we shaved o ff
.
1 5
o urn
En t
S oc
: 218
191 1
2 oo
and in each cell a hopper bearing an egg in precisely

the same p osition as be fo re One o f these eggs hatched
two days late r S ep tember 1 7 ; by the next day it w a s de
The other egg w a s p robably inj ured by
v el o p i n g rap idly
the crowding o f t oo many hoppers in a smal l tin box: Two
o f the hoppers d ied thre e days late r and one d ied a fter s ix
days The larva w hich hatched devoured its food speedily
leaving only the hardest parts of legs and integument and
i t grew w ith astoni shing rap idity Jus t nin e days a fter
the d iscovery o f the egg th e larva began to sp in I t made
a small a rea o f brown s ilken carpet under itsel f but ha d
di fculty in forming a cocoon in the small tin box which
was qu ite d i ffe rent from its natural earthen cell We tr i ed
to shap e a l i ttle nest o f cotton around it on wh i ch it m i ght
ge t a hold but i t could not adap t itsel f to i ts strange sur
roundings and died i n the attempt at cocoon making
Although we had l earned by thi s time j ust what to look
fo r w e succeede d in nding only one more o f these nests
during the summer o f 1 9 1 5 The characteristics o f th is nest
and its surroundings were in every w ay identical w ith those
j ust described I t was d iscovered only four days later than
the others and the larva on o n e o f the hoppers appeared
to b e about a day old
Anothe r season s observations were unsuccess ful i n clear
ing up the mystery o f the maker o f these nests s o we give
forth thi s m ea g e r record w hich really i s no more than to
make known the fact that th ere i s a ground digger p rob
ably a wasp which makes hori zontal tw in cells and store s
them w ith one locust each
.
'
C hl ori o n
c y cm e um
D a hlb
[ S A Rohwe r ]
.
We hereby enter our p rotest again st C Ma ri on c y an eum :

we spent ove r fty hours during o n e summer i ntently
201
watching and follow ing members o f thi s species foraging

without once seeing them do anything more than hunt O ne
could seldom spend even a l ittle while i n th e eld adj o in
ing the brick yard w ithout seeing one or more o f these blue
Chl ori om foraging among the cinders but w ithout getting
anything It seems p robabl e that they detect their p rey
o nly by the sense o f si ght
that they prowl about i n the i r
characteri stic manne r enteri ng holes and crevices for the
purpose o f nding p rey and that they are no t attracted
to certai n holes by othe r powers such as scent hearing
etc ; else they would n o t waste such a vast amount o f
time on hopelessly fruitless a rea s The p rey o f these wasps
is crickets and the hab itat o f crickets i s neve r in beds o f
cinde rs some o f them freshly dumped and still hot Yet
thes e wasps go on prowl ing about every accessible crevice
and c ontinue searching long a fte r one s patience and en
durance have bee n exhausted Sometimes they enter the
same hole many times wi th out nding anyt hing Thi s would
lead one to conclude that they hunt more o r less stup idly and
that e ach detai l o f thei r actions i s devo i d o f psycho logical
s igni cance ; po ssi bly they are attracted to the cinders by the
large number o f i nv iting crann ies and not by any i ndication
o f the presence o f cricket s
T w o y ear s ago an enti re a fternoon w a s spent i n watch ing
the similar behav io r o f these wasps at the f oot o f a straw
stack Although they did not to our knowl edge have the
good fo rtune to capt u re any c rickets they we re m o re j usti
ed in seeking them unde r w heat straw than in dry cinders
One warm morn ing i n Kansas we found a C cy a n eum in
a sandy plowed eld hurrying homeward w i th he r cricke t
A
robably
a
Serv
N
Caudell
She
l
u
b
r
e
v
i
a
m
s
l
a
G
r
s
b
[
]
p
y
travel led briskly he r cricket apparently no burden and lai d
i t down only once fo r a moment She carried it head fo r
ward unde r her body and at one time ew w ith it for about
,
2 02
three i nches S he m o ved in a man ner that i ndicated that

s he kn ew where she w a s going and
rtain
are
not
ce
e
w
e
t
y
that s he w a s go ing anywhere in particular
She went
straight across the eld f o r th irty feet until s he arr i ved at a
de ep furr o w down the s ide ; then s he turn ed at a ri ght angle
and followed di rectly down thi s for twenty feet more and
S he may have used i t w ithout
p lunged into a mol e hole
hesitat i on s imply because i t happ ened to be in her path but
s he travelle d w ith all the p reci sion and famil iarity o f a
man walking home from hi s o fce and turning in at hi s
own doo r S he rema i ned i n th e hole n i ne m inutes then
em erged hast i ly and soared o ff to t he woods agai n We tri ed
to dig out the burrow but ou r searches i n the loose sand
were all in va in
While i n thi s i nst a nce w e found C c y an eum dragging
he r cr i cket across the eld this method seems not to be
c onstant H unger ford and W ill iam s write that in G reely
County Kansas one was observed cl i mb ing a clay bank
carrying a mature female o f C en th op hi lus evidently str i v i ng
to reach an altitude su fcient to enabl e her to y to her nest
and the P eckhams nd that one a fte r runn i ng a l i ttle w a y
arose and ew l ightly for about e ighteen feet t o a hole on
the bare h ills ide H ere too P eckham s wasp di ffered from
ours i n that i t had its hole i n readiness on the sur face wh ile
o urs i f s he had a hole had i t c oncealed i n a rodent burrow
The P eckhams i llustrate a ve ry p retty burrow o f this wa sp
in which one egg wa s deposited am id seven crickets Gry ll us
From thei r later observations they nd
a b b r ev i a tus L i nn
she does not make a n ew nest for each egg but she p rovi
s ions a numb er o f cell s leading from one gallery They
record a very i nteresting i ncident wh ich gives on e the i m
.
'
16
17
16
1 7
Ent
N ew s
B ull W i s
.
2 3 : 24 7
912
N at H ist S oc
.
1 :
85
1 900
2 04
ing the wasp qui ckly ew out the door and escaped A few
nights later at twi light one o f thi s species p os s ibly the same
indiv idual but without her burden w a s yi ng about the i n
s ide wall o f the barn Eventually s he settled i n one o f the
hole s in the wall Th inking that thi s w a s he r nest and hop
ing to get more o f he r story w e di d not take her then B ut
on the morrow we so rrow fully learned that th is w a s o nly he r
s l eeping quarters and n o t he r ne st a t all

On e more eeting
gl imp se o f her som e days later carrying a large wheat straw
only tantal iz ed us and whetted our curios ity S o when
many days a fter the tw i ns came runn i ng to the house shout
ing I t s there ! It s ther e ! the wasp w i th a stick i n its

mouth
the fam ily b roke i n d iso rder and ran to the spot
Like the populace rushing forth cran i ng the i r necks sk y
ward to view the rst mode rn a i rcra ft to appear above a
village so w e ran h ithe r and th ither watching the m an euv res
ove rhead o f our grass carrie r wasp S he wheeled and ci rcled
about and nally landed at her burrow i n a log in the corn
crib I t was an ol d carpenter bee hol e completely stu ffed
w ith gra ss I t seemed that the straws had all been drawn i n
by the i r m i ddle so the en ds p rotruded f rom the orice in
a b room l ike tu ft We removed thi s mater i al and found
noth i ng but grass
H oweve r w e could not reach to the
bottom o f th e burrow w ith the forc eps
Th i s nest was the cou nterpa rt o f seve ral wh i ch w e had
seen about that time All o f them seemed to be ol d bee or
be etle burrows i n log s or t im b e rs an d al l we re s im il arly
lled w i th th i s material ti ghtly cramme d into the hole
These nests d id not appea r until nea r the end o f our soj ou rn
i n the country s o w e d id not learn rs t hand the us e o r
purpos e o f these straw structures so l ike a bi rd s n est
O n th e morn ing o f our last day w ith them w e sa w one
out gathering her gra sses S he al ighted on a lea fless branch
,
2 05
o f a plant and tried he r j aws on it but apparently it was too

tough She turned to a blade of green grass which bent
and s w ay ed un d er he r weight when s he al ighted on it AS
I expected to s ee he r
s he bit the blade o ff w ith her j aws
tumble when i t severed s ince s he w a s saw ing the branch
on wh ich s he w a s s itting B ut while the cut portion w a s
hanging by a mere thread she spread her w i ngs and ew
away b reaking it o ff as s he ew On her next trip s he
ew far away and returned a fte r fou r m inutes w ith a long
timothy stem protruding four inches f rom he r b o dy These
wasps always carry thei r grasses under the ir bodies grasp ing
the anterio r end in thei r mandibles and al ight and enter the
hol e w ithout shi fting thei r burden When w e v isited them
on Octobe r 3 thre e w ere at work all t he f orenoon busi ly
carrying dry grass into thei r chose n b ee-holes and even at
m
:
a
a
s
p
when
i
t
quite
cloudy
an
d
di
smal
one
w
s
w
4 45
stil l fai th ful to her task Thus the season ended w i thout
our having ascertained whethe r they use thi s mater i al for
bedding fo r food or a s a plug to close the ori ce
We a re not alone in thi s observation fo r P ackard nds

the same inse ct unde r the name S p hex tib i alis S t Fargea u
behaving somewhat s imilarly H e s ays M r J Angus who
reared thi s species sent h im a larva in a cavity previously
tunnelled by Xy l o c op a v i rg i n i c a in a p i ne board The hole
w a s s ix inche s long and the oval cyl indrical cocoons were
packed loosely s ide by side where there w a s room o r one a
l ittle i n advance o f the othe r The inte rstices were lled
w ith bits o f rope which had perhaps been bitten up by the
wasp itsel f The end o f the cell was lled fo r a distance o f
two inches w ith a coarse sedge arranged in layers a s i f
rammed i n l ike gun wadding The cocoons are 8 0 to 9 0
hundredths o f an inch long oval lanceolate somewhat like
,
'
20
G ui d e t o the S tudy
of
I n s ec t s p
,
68
1 889
2 06
those o f P o mp i lus They consist o f two l aye rs the outer

very thin and the inne r parchment l ike The larvae hib er
nate and turn to pupae i n the sp ring appearin g i n summer
and autumn
A s imilar typ e o f behav i or has been reported by Dav id
s on fo r a s ister species
unde r the name o f S p hex el ega m :
Fo r nestin g s i tes they pre fe r the larger stems rst gnaw

i ng thf o ugh the thin part i t i ons oppo s ite the lea f i nse rt i ons
wh ich naturally div i d e an otherwi s e ho llo w s tem
The
parent wasp rst packs the bottom o f thi s tub e w i th very
ne grass like be rs wh i ch on i nvestigation p rove to be
ne st rips o f the loos ely b rous bark o f Audib er ti a p oly
s tac ha
f rom one fourth i nch to one and one fourth i nches
i n length O n thi s i s laid the larval f oo d supply wh i ch
consi sts on an average o f from seven to eight tree cri ckets
The egg i s lai d on th e b reast o f one o f the crickets a wa d
o f bark be r i s placed on the top wh i ch fo rms the base o f
the next cell et c
A c opiou s wad som etimes thr e e or
four i nches i n dept h protects th e topmost cel l

,
'
'
The prey o f Amm o bi a
b ridw elli
Fe rn a ld [ S A Ro hwe r ] :
.
L ate one August a ft ernoon a t Mer amec H i ghl ands M i s

souri th i s wasp Wa s seen entering he r burrow i n the so ft
loose earth under a n open shed S ince it wa s impossible
fo r us to rema i n there w e captured the wasp and dug up
the nest I t w a s well provi sioned w ith the l ong a n ten n a ed
O rthopte ra C o m p ton o tus c aroli n em is G erst [ A N Cau
d ell ]
They w ere e ight i n numbe r three m
ales and ve
females The earth w a s so loose that w e co uld get no i dea
o f the shape o f the burrow
,
'
2 08
i nto the ai r where it made a l ittle pu ff o f d ust a s i t bl ew

away in the breeze
S he digs he r hole stra ight down fo r an inch or an inch and
a hal f then fashions an oval hori zontal pocket or chamber
abou t three fourths inch lo n g an d one hal f i nc h hi gh ( g
She works without e i the r haste o r hesitat ion but w ith
4
p recis i on Wh en at length the last mouth ful o f d irt has b e en
carried out and s he feel s that the burrow i s re a dy fo r us e
S he usually nds
s he comes out and hunts fo r a l ittl e clod
someth ing suitabl e w ithi n a few inches o f the hole but she
i s very exacting in her selection and w ill not be sati s ed
w ith anyth ing which fall s sh o rt o f her spec i ed requ ire
ments even though s he must seek fo r some tim e Thi s
clod must be j ust large enough barely to cove r the m outh o f
,
FI G
45
T he b ur ro
of
S p hex p i c tip en n is
N atura l s i z e
the hole or to t i nto i t l ike a pl ug but it must not fal l into

the burrow We have seen her use a b it o f stem o f j ust the
right si ze Th i s she places care fully ove r the entrance to
close i t to intrude rs during he r absence Sometimes s he
throws one o r two k i cks o f loose dirt ove r and arou nd the
spot mo re completely to disgui se the place Then s he goes
o ff on the wing i n l ow i g ht over the grass an d weed s i n
uest
o
f
prey
q
,
S AND LOVI N G AM M O P H ILA
T HE
2 09
The duration o f her absence varies Sometimes she meets

with ea rly succes s in her foraging and returns w ith her
cate rpi llar i n an hour ; sometimes s he i s out a s long as s ix
or eight hours presumably in he r search for prey We have
never been so fortunate as to w itn es s this part o f her work
We have seen he r out fora ging i n the eld to be sure
h0p p in g from lea f to lea f or ying over the grass scanning
the weeds a s s he lightly passes over them but w e have never
seen her at the critical moment o f the capture o f her p rey
B ut soone r or later s he comes trudging home w ith her
burden which o ften i s a grey cut worm At this part o f
he r task s he i s natu rally more cons p icuous and more easily
di sco vered than at an y o ther tim e We must not be sur
pri sed that she sometimes has some d ii c ulty i n progressing
with her load ; rathe r w e must be astonished that she can
move w ith i t at all The caterpillar i s very o ften four times
he r own weight or mo re yet she brings it through grass
and roo ts cl od s and gutters We have never seen an Am
m op hi la give up a caterp i llar because i t w a s too heavy fo r
her Sometime s howeve r the prey i s so small that s he can
drag i t w ith ease or even l i ft i t clear o f the ground and
make rapid p rogres s S he p ractically always carries it in
the same manner ; s he turns it o n i ts back s o the smoo th
rounded back w il l sl ide on th e ground l ike a sled runner ;
then s he stands astride it grasps the skin at the s ides o f
its throat until it i s drawn to l o ok l ike a tight c o llar
around the cate rp illar s neck l i fts the anterior end a l ittle
so only t he rounded dorsum wi ll touch the ground or o b
Once or twice we
s ta c l e s and away she goes on all six feet
h a ve seen h er gras p i t by the middle segment instead o f by
the thr o at and l i ft it clear o f the ground ( g
On o n e
occasion when a wasp was i n grave di fculties in getting
her caterp illar through a tangl e o f grass s he managed it
very ingeniously by li fting one end o f the burden and p rop
.
'
'
AFI ELD
WA S P STUDI E S
210
p ing i t almost vertical ly in the fork o f the g rass then l i ft

ing the other end to a slightly highe r po int Thus l i fting
,
FI G
46
S p hex p i c tip en n i s
c a r ry i n
s m a l l c a t e rp
ill a r
N a tura l
si
ze
the ends o f the caterp illar alternately s he h oisted it out o f

the chasm in th e grass thic k et in to wh ich i t had fallen
exactly as a m an working alone woul d ho ist a heavy tim
ber l ittle by l ittle Ou r A p i c tip enn is al ways walks forward
w ith her p rey beneath her as d id the Amm op hil a : oh
served by the P eckham s i n contradiction to Westwood w ho
says that Am m op hila when she has captured he r p rey
walks backward
I n thi s po i nt o f he r behav ior w e s ee the grossest erro r
i n economy comm itted by Am m op hila S he spends an
enormous amount o f energy and time i n trundl ing her
heavy caterp illar f rom a cons iderable distance w e know
not how fa r and then d ragging i t al l over the region i n
the v i cin ity o f her h ole wh il e locating th e burrow Since
s he uses but one ca terp illar s he might save all thi s needles s
waste o f time and ene rgy by do ing as P ri on on yx do es
d igging the hole conven iently near to the p rey a fter it has
been captured The topography about the nest i s u sually
very sim ilar i n nature to the env i ronment o f the p rey and
s he seems not to be ve ry particular about the location
To
be sure she has a p re ference for beaten paths and sunny
bal d areas but she as o ften uses any l ittl e bare spot b e
t w een t he weeds and grasses fo r he r nesting place So why
shoul d such a stup i d hab it p e rs ist entail ing frequent loss
danger and exp enditure o f e ffort while al l o f her other
methods o f work have become s o rened and highly devel
?
oped
The only explanation that w e have to o ffe r is that
,
'
212
that but each time when i t would not go in she woul d lay
i t asi de and di g the hol e deeper or at least enlarge the cham
be r at the bottom At last i n another attempt s he dragged
i t i n a fte r he r but i t becam e lodged wh en only one e ighth
o f i t wa s p rotruding hence she coul d ne ithe r push it o ut

n o r hersel f escape
We waited for a time and sa w that the
situation only meant death to her s o w e cam e to her ass ist
h
!
ance
Thus
our
human
ity
I
f
had
been
alone
w
ith
e
s
(
Nature s he would not have le ft any posterit y perhap s to p er
e
t
u
a
t
e
he
r
stupidity
!
We
pulled
i
t
out
w
ith
f
orceps
but
p
)
already the D ipterous paras ites wh ich had been lurking
nearby had d o ne the i r deadly wo rk fo r a hal f dozen squi rm
ing larva e we re at the neck o f the caterp illar As i t was
pulled out some o f these were brushed o ff and f ell down
into the hole S he showed neither ange r n or surpr is e at
the ass istance that had been rendered to her but she a p
r
e
d
satis
ed
w
ith
the
dispos
ition
o
f
her
p
roperty
S
he
a
e
p
came out and picked up a la rge rm pellet to o k i t down
into the hole w ith he r and remained quite a while brought
i t up fo r a moment then returned w i th i t and remained
ins ide fo r another per i od Then she cam e out once more
threw th i s p elle t aside and gathered a hal f dozen other
smalle r ones wh ich s he too k down into the burrow Oh i f
w e but could have seen what she w a s do ing w ith them down
there i n her dark chambe r ! D id s he use the rst large one
?
to kil l the paras ite s which in fested her nest
O r did s he
instinctively feel that i t wa s now time to bring i n material
?
fo r the partition o r p lug at the entrance to the chamber
H ardly the latter because whe n S he emerged s he promptly
d ragged he r caterp illar in again as i f nothing had happened
Thi s time s he succeede d i n getting i t a trie lower so the
rear end was level with the sur face o f the earth ; then she
seemed to encounte r the same di fculty a s b e fore Five
m inutes elapsed and s he could move it n either down nor up
,
S AN D LOVI N G AM M O P H ILA
T HE
21
Un fo rtunately w e had to go then so we dug up the nest

We watched care fully for the tiny D iptera larvae but i f
they were there they were lost in the digging
Two minutes i s all the time the wasp requi res for adj ust
ing her p rey and ovipositing and sometimes she i s through
in less time ( Figure 4 7 shows the position o f the egg on the
caterp illar ) Then begins the task o f lling the hole and i n
this s he works i n a most calm and purpose ful manner di s
turb ed o f course i f one crowds t oo nea r but w ith no s i lly
,
FI G
47
P o s i ti o n
eg g
on
c a t e rp
i lla r
N a tura l
si
ze
nervousness S he eme rges head rst and immediately gets

some small clods usually about th ree and takes them down
to the bottom We suspect that she arrange s these to serve as
a pa rtition between the chambe r and the burrow to prevent
the l o ose soil from being packed d o wn around the cate r
pilla r or crowded upo n the delicate eg g Then she j umps
out n imbly and scratches and kicks the loos e di rt nearby
into the hol e goes in and stands on he r head pounding and
packing i t down With each ram o f the head s he em its
a shrill buzz ing sou nd How th is buzz i s made we do not
know nor do w e know whether i t i s an expression o f pleas
ure or o f lab or or a mechanical part o f the e ffort Thus she
brushes in and packs down several layers o f l o ose di rt until
the hole i s lled almost to the to p
I f she uses all the
loose dirt lying nearby s he i s not i n the least di sconcerted
but simply b ites s ome more loose w ith he r mandibl es f rom
the sur face o f the gr o und and uses that When the de
pression i s yet about one fourth i nch deep she f etches a
large rm c l od sometimes as m uc h as ve times a s large
At rst it
as he r o w n head and throws it into the hol e
.
21
appears that s he i s using th i s method a s a qu ick w a y to

ll up he r hole B ut no sh e settles hersel f for work again
grasp s the clod rmly i n her m andibles and fo r a m inute
o r s o s he rubs and gri nds i t ercely against the bottom o f
the dep ression w ith a rotary or s ide to side mot i on unt i l
unde r the p ressure and wear th e cl od crumbles and i s worn
to dust an d b e comes part o f the compact ll ing She then
turns round and rou nd on top o f the hole sweep i ng to the
center the d irt w h ich has been pushed out at the edges o f the
sauce r shaped dep ression fetches another clod from some
whe re w ith in a di stance o f two o r three feet from the hole
and repeats the per fo rmance Thus s he uses sometimes as
many as ten o r twelve clods grinding beating an d b iting
them to p ieces unti l the top o f the ll i s level wi th the
ground I n th i s w a y s he works the ll down much tighte r
for w e hav e o ften seen her use several pellets and pound
them i n a fte r the hol e app eared level full
I n her next selection s he seem s to be mo re particular
She goes here and yonde r paus ing at clods and t i ny p ebbles
sometimes li fting them o r tu rn ing them over When nally
s he nds one which suits her fancy usually a pebble a l ittle
larger than her head but sometimes an unusually hard clod
o r bit o f w ood s he br i ngs i t i n her mandibles and grasp ing
it rmly s he rubs pounds and hammers down the d i rt on
the top o f the hol e until al l trace s o f the ll are obl i terated
When she has n ished w e ourselve s canno t di s ce rn the spot
H er task so skill fully do ne i s now at an end ; s he th rows
her tool aside a few inches and its away w i th an utterly
carele ss ai r as i f s he had f orgotten all interest w hatso ever
i n th is place and quite probably she has It i s i nteresting

t o note that s he cannot b e persuaded to use thi s tool be fo re
the p recise time fo r i t Once w e tossed he r a t i ny pebble
while she wa s yet busy grinding to pieces her cl od s w ith a
pestle and mortar motion but she only took i t w i thout ado
.
21
W A S P S TU D I E S
A F I EL D
that one might suspect that the wasp were here i m p ro

vi sing a tool w ith which to tamp down the sand
Considering t hat o ur tool using insect desc ri be d above
would h a ve n o thing to do w i th the pebbl e until a time
w hen i t w a s needed for tamp ing and in vi ew o f what i s to
follow a s the result o f very close observat i on by the tw o o f
us on anothe r p i c tip en n is m o ther cl osing her nest w e do
think that they actually have highly plast i c intellectual
powers Mor gan says } in speaking o f the interesting oh
servation o f the tool us i ng Am m op hi la o f the P eckhams :
Here w e have intel ligent behavio r ri s ing to a l evel to which

some would apply the term rational for the act may be held
to a fford evidence o f the perception o f th e relati on o f the
means employe d to an end to be attained and some general
conce p t i on o f pu rpo se
B ut H olmes tak es except i on to
this i nterp retation o f Morgan by say i ng : B ut i n estim a t

ing the p sych i c aspect o f the per formance w e must bea r i n
m ind that th e act i s o n e wh ich bo rders closely upo n the
normal i n stinctive behav io r o f the insect Th e seizure o f
pebbles in the mandibles and the packing i n o f d irt are parts
o f the i nstinctive process o f ll ing the hol e The wasp com
b i nes tw o features o f its hol e llin g instinct in a rather un
usual w ay Does she really perce ive the relation o f means
to end ? I am not sure that s he does

Wh et her Morgan s
o r H olmes i nterpre tation i s the correct o n e w e ho p e the
detai ls given belo w w i ll co nvinc e the reader In this c on
n ec t i on i t may not be am iss to relate exactly the conduct o f
anothe r o f thes e wasps through thi s part o f her task
The wasp when di scove red w a s car rying her caterp illa r
I t w a s a smaller wor m than usual s o small that she car

ried i t in her mandible s by grasp ing it in its m iddle and
l i fti ng i t clear o ff the ground so s he could us e al l her legs
f reely fo r p rogressi on S he made a pretty s ight carrying
so
H olm es T he E v o luti o n
,
of
A n i m al I n tel l i
en c e,
2 04
191 1
S AN D LOVI N G AM M O P H ILA
T HE
21
her prey in thi s new fashi on ed w a y A fter carrying it for

ten feet she dropped i t and started to remove the li d o ff
her ne st In our eagerness to s ee all one o f us pressed too
close and she ew away a short distance in alarm In a
moment she walked back to the cut worm exam ined it and
gave i t a reproving stin g on i ts unde r s ide near the p r o legs
but w e could not distingu ish which pair S he then strad
d led and sei zed it in the manne r al ready described and car
r ied i t o ff
making a very pe r fect ci rcl e about thre e feet in
di ameter and d roppe d i t at the identical spot from where
a moment be f ore she had indign antly taken i t A strange
w ay th is o f re sponding to disturbance !
S he then gave it
two i mpressi ons about the head w i th the mandibles then
passed the tip o f her a b domen on its unde r s ide as i t lay
ventral downward and felt o r rathe r p robed in a hal f

do z en places sl iding he r abdomen cont i nuously backward
unti l s he reach ed a spot j ust i n the rear o f the la st pai r o f
pro legs ; the re s he thrust in he r sting and let it rema i n for
several s econ d s We had thought at rst when s he fel t the
integument w ith her abdomen at a numbe r o f places that
she w a s stinging the caterp illar but when w e s a w the w a y
s he del iberately thrust i n her sting and let it qui etly remain
there for some time we deci ded that these fo rme r touches
had been only o f a t esting o r p robing nature S he then re
sumed her w ork where she had le ft o ff and opened her hole
by removing one m o uth ful o f clods She darted in and re
appeared almost instante r carri ed in her cherished prey
came out again and imme
remained ins ide t w o minutes
Whi le she w a s ins ide w e
d ia tely began to ll the hol e
took the oppo rtun ity o f observing the s ite closely There
w a s very littl e loose di rt or dust a b out the hol e ; all that had
been taken from the hole had been carried ve inches away
across a tu ft o f grass and piled up neatly in sp ite o f the

di fcult i e s of always clambering through the tangle o f grass
-
'
21
WAS P S TUDI E S AFI E LD
stems S he came out head rst and carri ed in one at a time

three clods o f earth ; these w e suppose s he arranged to for m
a plug in the angl e o f th e L shaped nest A fte r that fo r

a few seconds s he kicked loose dust i nto the burrow
S he
also k icked a straw s ix inche s in length over the hol e ; thi s
s he carried a few inch es away
and later when it got i n
her way a second time s he carr ied i t i n her mandibles and
ew w ith i t for so me d i stance and cast i t a way S he kicked
i n more loose earth and a fter each portion went in to ram
or pack i t down humming i n a discontinuous w ay that i s

i t sounded as i f w ith each ram o f he r head she noti ed us
w ith a sharp buzz o f l ike duration
Thus s he used all the loos e d irt that lay near at hand ; the
region w a s hard and clean There wa s yet about one fourth
inch o f the burrow to be lled when she came w ith a hard
clod about ve times a s large as he r own head threw i t i nto
the hol e and went i n hersel f as far as p os s ibl e We thought
thi s woul d be an easy way t o ll up the burrow and were
surp r i sed to s ee he r grasp i t i n her mandibles and continu
o us ly for three m inutes p ress d o wn the earth in the d ep res
s ion w ith th i s tool rubb ing w ith i t i n a circular mot i on
B ut ou r surp ri se w a s greate r when w e saw he r throw out
thi s hard clod wh en s he had t a m p ed the dirt d o wn s uf
A
c i en tly ; the easy way o f ll ing a hole w a s not he r w a y
much smaller clo d w a s then thrown i n and beaten to dust
w ith her head and face Th is w a s plainly to be s een and
thi s later clod must have been i ntended for thi s purpo se since
the mandibles and head were us ed to break i t up and no
attempt was made t o sw ing i t i n a ci rcular motion I t i s
quite l ikely that th i s small one could also have served as a
to o l i f she had wanted to use i t to that pu rpose and we fe el
sure al so that the former one could have been b itten and
pounded to p ieces w ith her stout mand ibles had s he wi shed
to d o s o B ut s he had done the nece ssary p ounding ; lle r
.
'
22o
I
I
x
\
5'
J
t
I
'
"
J
t
K
R
>\
'
b
\
J
a
FI G
48
D i a g ra m
th e l a n d m a r
to
ab ou
sho
h er
w th
n es t
w ay
one
m o the r 5
p i c tip en n i s
s tudi e d
away from thei r burr o ws l eavi ng suddenly w i thout gett i ng

any landmarks they have grave di fculties in return ing to
the i r nests O ccas i onally on e departs on the w ing f rom he r
incompleted burrow w ithout any ap p earance o f exam ining
the region M o re commonly howeve r they go around the
covered n est in w idening ci rcles rst walking near th e hole
then running h op ping an d at last ying i n l arge r ci rcles
and nally soar i ng o ff t o th e elds i n quest o f prey One
wasp which we stealthily watched seemed unusually p recise
i n her manne r o f examining the locat i on be fore departing
,
TH E SAN D LOVI N G AM M O P H ILA

-
hunting grounds
22 1
for the
The hole w a s nished and she
had brought three p ieces o f di rt and ca re fully covered i t
S he ew to a lea f a few inches away From this perch
s he descended thre e times t o do some unnecessary scratching
over the hole B y thes e m aneuvers s he m ay have b ee n get
ting her landmarks ( s ee g 4 8 ) i n fact we sus p ect that

s he w a s f o r s he walked al l o ver this small S o la n um ( S i n
the gure ) which o verhung her burrow ( N ) ex am m i n g I t
all the whi le an d as she came to the lea f di rectly above he r
nest s he usually pee red d own and s om etimes desc ended to do
some scratching then retu rn ed to continue he r examination
o f the weed Then s he turned he r attention to a small tu ft
o f gras s ( G ) the o bj ect next i n nearness to her nest and
scrutin ized i t A fte r thi s m inute exam i nation sh e dropped
down upon the earth and walked in an almost per fect circle
around the nest not all at once but i n arcs and a t th e ter
mination o f each arc she would waltz abo ut in t i ny ci rcles
sometime s one wh i rl and so metimes two o r even three times
around and then p roceed w i th the next arc without o nc e
becom ing con fused in he r d i rection s or m issin g th e lines o f
he r well de n ed ci rcl e The l ittl e waltz i ng whi rl s we re a l
most always from le ft to right but once she ci rcled f ro m
right to le ft The accompanying crude diagram ( g 4 8 )
will ai d i n making clear thi s rema rkable p er formance While
w e had no measure o f the len gth o f the arcs it seemed to
us they were wonder fully uni fo rm i n size This prec i s ion
i s only i n harmony w ith th e othe r in stances o f ne physical
j udgment wh ich w e s o o ften s ee in the conduct o f th is wasp
At the conclusion o f the rst ci rcl e s he c ommenced a second
o f increasing rad ius but a s she progressed she got to run
ning and hopp ing and soon ew away to the vegetation
beyond A moment late r w e s aw her climbin g up a rag
weed in the r o l e o f a forager
She le ft at I O o cl ock an d w e could not wa i t for her return
.
WA S P STUDI E S AFIELD
222
but at 4 :30 p m w e exam ined the nest and were surprised

to nd it j ust a s we had l e ft it We removed the l ittl e lumps
o f clay w ith wh ich s he had covered it but the ho le w a s
empty s o w e replaced them ; no prey a fte r s ix and one hal f
hours ! We b egan to wonder i f s he had gotten lost in spite
o f her accurate o r i entation
Early the next m orning w e again exam ine d the sp o t and
found i t tightly lled in A l ittl e care ful digging soon re
veal ed the cut worm w ith the del icate egg on its side
H ence s he must have return ed to the n est w i th the prey
som e time between 4 : 30 and sunset a fter an absence and
a search o f s ix hours o r more H e r local ity study served the
purpo se o f her accurate return al though many marks were
requ i red to guide our return
I n anothe r case th e burrow w a s ready fo r use and the
wa s p p roceeded t o close i t temporarily as described else
where w hile s he went foraging H e re her j udgment must
somehow have been am i ss fo r the rst two pellets which she
b rought w ere s o small that they fell down i nto the hol e ; but
the th i rd w a s large enough to rest on top n i cely l ike a l id
The n s he concealed the s p ot de ftly by adding some othe r
b its o f earth and a k i ck or two o f dust and departed for
green eld s S he did not tak e the usual ight o f orien ta
tion but began by searching 1 n the weeds nearby
S he
itte d o r hoppe d f rom plant to plant but whi le doing thi s
her path described a c i rcle around th e hole S uddenly s he
ew d i rectly away out o f our s ight Five m i nutes l ater s he
returned and continued scanning the weeds i n thi s region
and at one t ime even walked ove r her own nest and stopped
short o n her p il e o f pellets fo r a mom ent then ew away
In observing pi c tip en n is the fact i s dem o nstrated ag ain and
a gain that the insects w hen returnin g to the obj ect o f the i r
search are n ot guided b y any speci al sense but in all prob
abil ity as th e P eckhams Turne r and othe rs have shown for
.
224
w ith he r and sometime s l a ying it down At last s he seem ed

to stumble up o n i t and then turned back to get he r bur
den wh i ch w a s nea rby I n her con fus ion by th is time
howeve r s he pulled loos e from the earth another l i ttle clod
and used that a s a covering f o r he r burrow Thus it seems
that even a little scrap o f pape r bes ide the nest s o altered
the t o pography as to utte rly con fus e thi s wa s p
Another wh i ch l e ft l ikew i s e unde r p rovocation roamed
about i n t he reg i on o f he r prey fo r an hour wh ile w e waited
at a sa fe distance w ith the camera focused upon he r cater
pillar A fte r a ted iou s hour w e b oth l e ft i n d i sgust s he
w i thout her cate rp illar and we w i th o ut our p icture
I nstances o f thi s so rt only substant i ate th e place mem
o ry idea When s he suddenly l eave s her p rey in the m idst

o f her j o urney and has no opportun i ty o f studying the land
marks even though s he be i n the immediate v icin ity th i s
wasp i s unable or able only w ith great d if c uly to nd her
prey When s he does nd it a s s he occasionally does we b e
l i eve i t i s due merely to her stumbl ing upon i t by chance as
and i f the same element o f chance
s he scours the region
leads he r footstep s one i nch to the r i ght or l e ft o f i t i t i s a s
lost to he r as i f s he were a m i le away I t d oe s not seem that
scent o r ev en s i ght pl a ys a very g reat part at l ea st the
sight o f the o bj ect o f he r search although p robably s he

must de p e nd upon the v i su a l s ense i n establi sh i ng her pl ace
memories I t is barely poss i ble that th is m igh t have devel
oped a cond i t i on o f far s i ghtednes s
P i c tip en m s i s ve ry sel ect i n the choi ce and treatment o f
he r p rey We have neve r known he r to us e more than a
singl e i nd i vidual for th e p rov i s i o n ing o f a nest but these
are sleek fat and meaty usually o f the cut worm type Some
o f them have been identi ed fo r u s by D r B a rnes and Dr
M c D o n n o ugh a s Agros ti s C n ig rum and a H esperid la rva
Another species o ften u s ed i s identical
P holis ora c d tullus
.
'
SAN D LOVI N G AM M O P H ILA
T HE
22 5
w ith the i llustration in Holland s Moth B ook o f Ly c i a c og

Th is i s a G eometrid that i s known to be
n a ta ri an G uen e
protectively colo red and sti ffens out to resembl e a dead twig
when attacked yet somehow p i c tip en n is gets i t Anothe r
Lep ido pterous larva used by this wasp i s P ro dem a orn i tho
l
B
G
uen
Fracker
One
green
caterpi
llar
o
f
the
a
l
i
S
[
]
g
N o c tuida e [ S B Fracke r ] taken from p i c tip en ni s while I n
transportation l ived eight days
The imp ress ion exists that the cut worms used by Amm o
i
Alm
o
st
r
h
l
are
subterranean
ate
pillars
Kell
gg
says
:
c
o
a
p
all cut worms hide in cracks in the ground by day feeding
during the night and since cut w o rms are so o ften used
by p i c tip en n is i t would seem that s he would be obliged to
acqui re the subterranean hunting habits o f her Europe an
cousins
In fact Turner describes the behavior o f an
Ame rican Am m op hi la s p that digs in the ground for cater
p illars w i th which to stock the nest Thi s lea d s him to con
c l ud e that
some o f ou r Ame rican Am m op hi las resemble
those studied by Fabre in storing the i r nests w ith s ub ter
ra n ea n caterpillars
We have neve r been so fortunate a s to o bserve these
was p s at the m o ment o f capture o f th e p rey but on one
occasion w e s a w one whose behavio r w a s such as might
easily arouse suspicion
Thi s insect was running about
a smal l plot o f gro und in s o strange a manne r that our
interest w a s at once attracted It would take short ights
in the ai r j ust abo ve the ground and suddenly drop to the
ground w ith a sort o f thump landing on al l i ts legs as i f
sounding the ea rth H ere i t would walk slowly about the
spot for a few seco nds then wheel round i n anothe r short
ight and repeat the pe r fo rmance This i t continued to do
for twenty m inutes until at length i t seemed to decide upon
'
Am
I n s ec t s
P s y c he
: 1
402
191 1
905
226
a certain spot in the broad barren area whe re i t began to

dig It woul d stand squarely on its head and hum a
high p itched l ittle tune whil e at work I n digging the man
d ib l es and fo relegs were used to chop and break the dry earth
whil e the m iddl e legs were u sed to brush the dust away at
both s ides At rst there w a s n ot hin g to be brushed away
yet this pai r o f legs worked automatically and p rettily
There was no S ign o f the usual habi t o f carrying the ex c a
v a ted di rt away t o a p ile as i s always done when the burrow
i s being dug With the hind legs alon e t he wasp held rmly
to the ground I t l e ft the spot several times as i f i n doubt
o r di ssati s ed about so mething but returned
I n a few
minutes a n eat dep ress i on was made ; then w ithout p rovo
cation o r without wa rn ing o r ight o f orientation it arose
and ew fa r away never to return Was i t i n quest o f
prey which i n thi s instance i t di d not nd or are w e m is
j udging i t in thi s susp ici on ?
We have never found A p i c tip en n is to provi sion her nest
w ith mo re than one caterpi lla r although the P eckhams
found that A m a ri a stores two caterpillars i n each nest
Wi ll i ston nds that A y a rrow i Cr stores fou r or ve cat
says that the large A p r oc era cap

e rp ill a rs and H artman
ture and sto re but o ne immense green tomat o caterp illa r
wh ile the smaller specimens o f p r o c era bring i n three cater
p i llars Kellogg nds that an Am m op hila s p wh ich i n
habits the sal t marshes near Stan ford Univers ity uses fro m
ve to ten paralyzed inch worms f or each young wasp

According to S harp A lzirs u ta chooses a single larva
o f consi de rable s iz e belonging to o n e o f the nocturnal
1
Lep idoptera
Fabre nds that A juli i stocks each
,
10
'
'
B ull W i s G eo l
B ull Un iv T ex
7
8
Am
1
1 1
I n s ec t s
I n s ec t s
H un ti n
N at H i s t S u rv
Pt
p
2,
66
49 4
.
W a sp s
: I
9 05
2 : 25
III
pp
89 5
259,
419
91 7
89 8
WA S P
228
STUDI E S AFI E LD
wasp in he r long hop eles s struggle to reach he r burrow

The second o r fo raging wasp became aware o f its p resence
whe n she was withi n an i nch or two o f it

sni ffed a ll
aroun d w ith her antennae and at o nce located it She

d id not pounce upon the caterp illar w ith violence as we had
expected but paused and exam ined i t closely running to
and f ro eagerly exami nin g rst one end and then the other
alternately s everal times t hen pinched it i n a numbe r o f
places along the ventral su r face and bit it on the throat close
under its head seve ral times s o v iciously that w e thought
that s he w as delibe rately decapit a ting it Then s he rel ea sed
her grip and turned it over dorsum up mounted its back
and curved her abdomen de ftly around i t until s he planted
he r stin g be hin d the rst pai r o f pro legs The caterpi llar
whi ch had hereto fore la i n motionless drew up quivering
Then the wasp w ithdrew and inserted her sting behi nd the
second pai r administe ring a prolonged and most del iberate
sting and also behin d the thi rd and fourth fou r sl ow sure
stings Then s he l i fted and inqui sitively shoo k its head
and then shook its tail :1 b it a s i f to make sure that it w a s
r eally don e for turned i t ove r o n its back straddl ed i t and
w it h some d i fculty ( for it was an exceptionally large and
heavy caterp illar ) got started with it across the smoo th
ground S he grasped i t w ith a terrible grip o f her mand i
bles and part o f the t i me w i th her rst legs too
She
dragged i t i n the usual fash i on dorsal s ide down Then
followed the endless chase ; s he seemed now t o b e hunting
fo r he r nest in this local ity and s ometimes a p peared to be
preparing to dig as i f to uncover it but it may have been
destroyed o r sh e may have been badly mi staken i n her
loc ation
S he trailed he re there and yonde r s o metimes
several feet away and then came back to thi s vicini ty only
to wander o ff again She ran around s o much w ith that
caterp illar that we thought surely it would be worn to shreds
.
T H E S AN D LOVI N G AM M O P H ILA
229
be fore she woul d use it As time passed by her j ourneys

away fro m the original point became longer A thunder
storm was app roaching but neither the dark clouds nor the
rst patte ring drop s o f ra in drove her to des ist Thi s w a n
de ring and va in search had c o ntinued relentless ly for over
two hours unde r the August s un and the dark c lo uds ; and
we wonde r how much longer she wo u ld have continued it
had not a horde o f ro m
ping boys passed by and frightened
her away
Thus s he apparently accepted th is pro ffered caterpillar
and treated it as he r own The stinging certainly could not
have bee n done with more pr eci si on T he malax ati on i s
sai d to b e a regula r part o f Am m op hi la s treatment o f her
prey and occasionally i f a v ictim does not seem wholly
and completely submi ssive s he w ill pause i n her j ourney
and administe r some more p inching kneading and shaking
Again and aga in we have been asto ni shed a t th e remark
able tameness o f Am m op hi la when building provi sioning
or cl o sing her nest Th i s cannot be attributed a s we thought
at one time to de fective eye sight for when s he is al o ne
w ithout a caterp illar o r a nest to detain he r she i s very
evas ive and usual ly many attempts to approach her are
futile B ut when he r interest i s abso rbed i n home dutie s
she seem s sel f forget ful an d o n e c an u sually appro ach very
near to he r withou t he r g ivin g heed On several occas ions
when a wasp w a s in di fculties i n moving her caterpillar
we as s i sted her with her burde n w ith our force p s and
s he seemed to take i t all as a matte r o f co urse a s i f w e were
friendly collaborato rs Once in o rder to s ee how s he would
respond to teas ing we took hol d o f a cate rpilla r which an
Am m op hi la was bringing to he r hole
She would not re
linquish he r hold but gripped the o the r end braced he rs el f
with spread legs and pulled with all her m ight A tiny
tug o war ensued until we l i fted her clea r o ff the ground

.
WA S P STUDI E S AFIELD
2 3o
still clinging to he r treasure S everal times w e thus l i fted

the caterp illar and the cl inging wasp into m id a ir and each
time s he felt hersel f once more on rm earth s he again
dragged her prey ove r to the brink o f her hole On another
sunny S eptember morning the feminine partner in th ese
obse rvations was s itting u p on the ground bus ily w riting
up the notes o f the mo rning s observations when an Am m o
il
cam
e
and
began
i
ndustriously
digging
a
burrow
right
h
a
p
bes ide he r knee whil e another homeward bound w ith its
prey cl imbed up on he r dress and trundled its caterp illa
down the full len g th o f he r bare forearm pausing once and
laying the cut worm down on he r hand whil e it washed its
face and looked about inqui ringly then moving calmly on to
the burrow beyond
Fi nally one day we went so far a s to take the cate rp illar
away f rom a wasp and place i t in the hollow o f the hand
at on the ground p erhaps eight inch es away f rom her
The wasp came w ithout hes itation and got it ; in fact s he
betrayed less nervousness than w e fo r s he seemed to fear
nothing while we remembere d very v ivi dly how wasp s
o ften re sting caterp illars which m i sbehave and feared tha t
her a im might not always be accurate
Later w e removed the caterp illar from the mouth o f an
other wasp placed it in the bottom o f a two inch vial and
laid i t at upon the ground The wasp soo n found i t ew
i n circles j ust above the v ial and nally rested on top o f it
I t was ev ident
f or a t i me then agai n ew ab o ve it in ci rcles
that s he s aw her p r op erty but e ither s he did not nd the
opening or fea red to go so far i nto a myste rious place for
he r p rey We then placed her caterpi llar i n the neck o f the
vial whereupon s he promptly went into the bottl e turned
around to get her cust omary grip upon the caterp illar and
happily trundled it o ff hom eward
A fte r s he had g o ne
some distance w e agai n robbed he r and placed her treasure
.
WA S P STUD I E S A F I ELD
2 32
baseball diamond at rst rathe r ne rvously w ith twitching

w ings then calmly as i f her m ind were all on he r quiet hom e
dutie s Then w e s a w that she w a s carry i ng l ittl e cl ods
about as large as her head I t seemed that s he was trying to
lay them on top o f the hol e but they all w ent ins ide S he
did not h o wever drop them i n w ith a thum p but she went
i n w ith them and cam e o ut at once in search o f mo re She
walked all the time ( neve r ew ) and seemed to become
more calm and serious a s the work p rogressed S he walked
he re and the re sei zi ng every protuberance on the sur face
o f the hard packed earth that might serve her and giving
it a vigorou s erk to di slodge it P resently s he reversed her
act i on to ok out the clods one by on e and threw them a w a y
until s he had discarded seven clods o f the s ize o f her head ;
then s he brought in thei r place one large clod w h i ch w a s
about tw i ce the s ize o f the rst one s and o f precisely the size
needed S he la i d it over the opening and b rushed up a l ittle
loose d irt oh so nicely ove r and around it and departed

In taking her leave s he walked around the nest at rst , then
ci rcl ed around by short j umps gradually lengt hening into
one two and three foot ights then a continuous ight
in a w ider ci rcle and away s he s oared to the weeds We
can s ee no explanation o f thi s j uggling w ith the clods other
than that e i ther th i s w a s he r rst nest o r s he was not gi fted
w ith the usual i nstinct or skill i n j udgi ng exactly the r i ght
s ize o f cl od and she learned i t o nly by experimenting At a
later stage in the nest building clods are regularly taken
down into the burrow ; i s i t not possibl e that thi s female w ho
w a s probably possessed o f de fective instinct i n one phase
or anothe r had momentarily con fused this temporary closing
?
w ith the permanent clo sing
Afte r the caterp illar has been stored i n the chamber at the
bottom o f the burrow a plug o f cl o ds i s alway s placed in
the mouth o f the chamber to prevent the d ir t fall ing in
,
TH E S AND LOVI N G
A M M O P H ILA
2 33
aro und i t when the gallery is lled Th is too is tted i n

w ith pretty accuracy
Sometimes a singl e clod i s used
sometimes two or three form the partition I n one nest
which we dug out the mother had utilized a round pellet
o f rabbit s dung for thi s purpose I t tted the hole s o n icely
that it must have been forced i n with cons idera ble energy
Even in thei r ways o f meeting obstacles which chance to
come i n thei r w a y Amm op hi la: display more than the
average good sense They do not merely kick the o bj ects
back out o f thei r w a y tem p o ra ri ly b ut they p ick them up and
carry them aw ay or in the case o f straws or lea f fragments
they y wi th them to a sa fe place t o deposit them How
unl ike t hi s i s to the stupid P omp i lus m a rg i na tus which we
sa w digging in the bottom o f a l ittle funnel shaped d ep res
s ion w here a clod rolled i n upon i t and seriously h indered
i t S ix times the silly creature carried that clod up and lai d
it on the brink o f the depression from where as soon as
he r back w a s turn ed it r o lled down again and bumped he r
Thus A p i c tip en n is seems to be a ti dy and exact housekeeper
i n all details ; s he always carries out the dirt and p iles i t up
in a neat mound at a certain d istance f rom the hole and
i n the direction op p os ite the hori zontal chambe r at the
bottom I n one case which w e noticed the site o f the hole
was surrounded by grass but even thi s di fculty did not
de feat her ; she carried all her digg i ngs to the appo inted
spot even though she had to carry them through a tangled
tu ft o f grass which must have been to her a veritabl e j ungle
Another abnormal instance which caught our attention
was an unusually large nest I t was al ready about two
i nches deep a l ittle beyond normal depth when we found it
and the wasp wa s continu i ng the excavating There w ere
two piles o f di rt ; o n e the smaller was almost at the brink
o f the hol e and the other twelve inches away S he was
carryin g all he r di g gings to the more di stant pile all the time
.
WA S P S TU D I E S A F I ELD
2 34
watched her S he carried large loa ds and the so il was

sandy and crumbled readily ; s o s he bore each load with
b o th f ront legs around it S ome species o f wasp s reserve
n ear at hand j ust e n ough dirt t o ll up the channels o f their
burrows w hen they close them and throw away all that
c o mes from the deeper chambe r I t i s po ssible that this indi
v i dual showed a reversion to thi s typ e or w a s a sport j ust
develop ing it but we fancy that it i s more probabl e that she
was merely a f reak s ince when we dug up th e hole w e
found the hori zontal chamber much large r than usual The
caterpillar on e o f only medium s ize seemed quite lost in it
I t m ight be that the pile near th e ori ce had been accidentally
dropped when s he emerged from so great a depth I t may
seem to s ome that w e give an undue amount o f attention to
petty detail s o f th is orde r but w e want to record every po int
and search every detail wh ich m ight lead o r contribute to
knowl edge o f the ori g in o f new habits
The duration o f the period o f developmen t o f the larva o f
Am m op hi la or the t im e from the deposition o f the egg until
the pupation o f the larva i s brie f but somewhat variabl e
One usual ly nds the egg hatched on the second day unless
it has been depos ited late i n the day i n w hich event the l ittle
larva about ve m m i n length i s not present unti l two days
a fter oviposition B y the fth day o f its existence the
larva usually becomes large fat and white and the next day
or at most two days therea fte r i t sp ins its whi te s ilky c o
coon During the grow ing stages the larva s eems only to
suck the j uices o f its host ; but i f it i s hungry on the last day
o f i ts go rmandiz ing i t sometimes devours eve n the tough
w ithered carcass o f the caterpillar When i t has attained
its ful l growth and i s ready to spin i t announces the fact
by a peculi ar way o f w rithing t o and fro We have not yet
ascertained the real reason for these conto rt i ons
How
ever i t i s inte resting to note that Amm op hi la always pupates
we
W A S P STU D I E S A F I EL D
2 36
caterpillars ies tent caterp illars sp iders taken f rom mud

nest s or pupa e o f Tryp oxy l on
Am m op hi la l ike all the others i s haunted by persi stent
parasites One l ittl e D ipteron o ften follows her l ike a bad
conscience pe rsi sting unwaveringly for two hours o r more
at a time as s he drags he r caterp illar here and yonde r in
searc h o f her hole poi s in g in m i d a i r always abou t twelve
or fourteen inches beh ind he r advancin g retreating tur n
ing w ith every movement o f her leader a s i f rmly attached
to her in som e invi s ible w a y I f p i c tip en n i s lays her cater
p illar do w n and leaves it fo r a time while s he searches
thi s paras ite w ill s i t down at her usual s a fe distance and
watch i t unti l th e ow ner comes b ack and claims i t L ike
parasites o f other wasps it seems to have no interest i n a
host le ft lying thus exposed but i s eage r about i t as soon
as it i s taken away with a p rospect o f be ing burie d p roperly
We are unp repared to make any statements regarding
the frequency o f paras itization but it i s not uncommon for
w e have several times dug up nests only to nd the cate r
p illar heav ily in fested w ith D ipterous o r Hymenopterous
larvae i nstead o f serving as nouri shment fo r Am m op hi la s
larva
The adults too have the i r enem ies We o nce rescued a
ff
whi
ch
being
carried
o
i
n
the
mouth
o
f
a
n
a
s
w
c
t
i
e
n
i
s
i
p
p
robber y P r o c tac cm thus m i lb er ti M acq an d M r G eorge
M G reene records that he found the adults stylopi zed by
-
1 5
E up a tho c er a
The adults hav e been seen feeding on the nectar o f the

dandelion and Robertson records them frequenting about
eight speci es o f owers
We h ave only onc e caught a gl impse o f the matin g o f
thi s species I n the dooryard o f a farm house one June
1 6
1 5
16
Ent
N ew s
28 :
94
T ra n s Ac a d S c i S t
.
1
.
91 7
Lo ui s 6
45 5
894
TH E S AND LOVI N G AM M O P H ILA
2 37
day we were watch ing an A p i c tip en n is taking her caterpil lar

home at a gall op ing speed S uddenly we knew not whence
a male swooped down upon her She was knock ed from her
caterpillar but at once resum ed he r work ; s he appe ared
much con fused and bothered by hi s persi stent i ntrusion but
she di d not pay him any particular heed To ou r surpri se
another mal e in great agitation dashed down and v iolently
bumped the rst one and for a few seconds fought for her
possession ; then i t departed A fter perhaps three m inutes
the rst male departed unce remoniously The female
showed no conce rn for either hi s presence o r hi s departure
but went ahead w ith her work a s though glad to be at peace
aga i n
.
S p hex
A
o
h
m
m
h
D
h
l
A
Ro
wer
i
a
r
o
a
l
b
c
e
ra
S
(
[
]
p
) p
.
A three m ile walk along a country road on S eptembe r 9

gave us many glimpses o f Am m op hi la p ro c era alon g the
w ay
We s oo n l earned that when w e s a w a membe r o f th is
speci es i n the sunny s p ots i n the road we would presently s ee
the second nearby They seem always to play and frol ic in
pai rs The two woul d whi rl about each other in the ai r
i n a play ful manne r then one would suddenly y in a straight
l ine along the wagon track i n the road with the other close
behind it The leader woul d then come to a re st o n the
r 0ad o r on a stone and the other would at once do the
same some di stance away Sometimes they would sit thus
ignoring one another for from three to ve minutes w hen
one would rise and hove r or p o i se ove r the othe r restin g
individual ; thi s seemed to be a signal to follow and so i t
usually did S ometimes ho weve r like play ful children they
c o uld not readily be coa x e d a way i n thi s m anner but the
l eader would ma ke a f e w circu its i n the ai r a few inche s
-
'
2 38
from the resti ng body Occasionally it p roved di fcult to

a rouse the playmate and sometimes i t w a s necessary to a d
m ini ster a few hard blows w ith the abdomen on the abdomen
o f the lo iterer to make i t wake up Thi s i s not mating Thi s
pe r form an ce o f dancing i n the ai r l es s than ve f eet f rom
the ground chas ing on e another on the w ing resting at
upon the ground near together has happened s o o ften in
our observations in various times and p laces that w e feel
con dent that th i s play i s a constant and characteri sti c hab it
o f A p roc era and not the antic s of a n i ndiv i dual or group
We are also su re that i t i s play an d not c ou rtshi p for
among the one hundred or mo re see n th a t day only two
were females
At Fox Creek on August 2 w e s a w d i fferent a spects o f
the mating o f A pr oc era o n the sandbar H e re there were
about a hundred male s and o nly three females The males
we re chasing on e anothe r an d y i ng ab o ut near the ground
so w i l dly and play fully that w e coul d not tell what they were
until we took them i n the net B ut the females behavior
was di fferent They woul d walk about slowly i n a smal l
space as i f under the p retext o f l o ok ing fo r something
S ometimes during a b rie f peri od o f th is a numbe r o f males
would at di ffe rent times sw o op down for an instant
a n eo us mating then y away again and j oi n the others i n
the dance seldom or never returning to her Thi s behav ior
i s som ewhat di fferent from the one c ase observed a few
years ago at Lake V i ew Kansas Three p r oc era two males
and a female cling i ng togethe r i n a mass were ying
slowly from plant to plant
At Moselle
m any large A p r oc era were yin g abo ut
in courtship no t on the hot sand but usually in the low w et
places near a stream They never rested long enough fo r
us to ascertai n the s peci es w ithout capturing them One
would follow anot h e r in a straight l i ne for a l ong distance ;
.
2 4o
At Allenton Mo July 1 1 1 9 1 6 one o f these wasps was
s earching for something hal f w ay up an almost vertical c l ay

bank ve feet high S oon s he di sc o vered he r big green
caterp illa r N a da za g i b b os a [ S B F ra c k er] which w a s lying
there ; she took i t up between he r legs and continued her
j ourney straight up thi s stee p bank w ith aston ish ing rap idity
S he hel d i t j ust a s A p i c tip en n is does w ith the dorsum o f
the caterp illar toward the ground although i n this case such
a pos ition seemed unnecessary si nce s he held the caterp illar
whi ch must have weighed about eight times as much as she
high enough f r o m the ground to make fri cti o n impos sible
I t was di fcult to f ollow he r among the weeds o n top o f the
bank I t i s doubt ful i f she knew where s he was go ing si nce
three times i n cours e o f he r rambl ing travel s s he cl imbed the
trunks o f tree s t o th e height o f three to v e f eet as i f s he
were getting her bearings ; then she woul d y down w ith
he r burden and continue he r j ourney P resently s he aban
d o n ed he r caterp i llar seeming unabl e to reach he r desti na
tion
When the caterp i lla r N ad a ta gi b b os a, was p icked up a fter
s he had di scarded it i t seemed unable to respond to stimul i
i n any but the last s egment o f the bod y The th ird day it
was stil l al i ve a nd responded w ith both anterior and p o s
te ri o r extremities
When one walks along the road and sees as described
above A p ro c era: y ahead o f one i n a straight l ine pause
a moment then y furthe r on etc o ne thinks that
this might se rve as a method for dissemination and
that wasp s continuing thi s for a number o f hours each day
for several days m ight soon be fa r from thei r fatherland
B ut a fter observing thei r ways care fully w e are su re that
they do not roam very fa r f rom home although they appear
to On that day i n particular w e follow ed many wasps
They would y along the road in a strai ght l ine and lure us
.
'
'
T H E S AND LOVI N G AM MO P H ILA
241
but since they coul d n ot outdistance u s w e soon found

that a fter they had travelled thus from three hundred to
ve hundr ed feet they i nvariably turned back I f w e con
t i n ued to foll ow them on the i r return we would arrive nearly
at and sometimes beyond the starting point where they
would usually disappear i n the weeds We could s ee no
femal es at work along the roadsides Even though the
mal es should mi grate along the highways thi s alone would
not be a factor in thei r di ssem ination while the females
remai n near the old hom e since the female alone decides the
hom e s ite The abundance o f males shows that they do not
die o ff as early as do the males o f B em b ix n ubi lip en ni s
We have been abl e to examine only t w o s p ecies o f Am m o
hil
a
i n the eld A p i c tip en n is and A p r oc era but w e feel
p
that there i s some l ittle evidence to i ndicate that A p i oti
n
n
i
e
A
A
i
s
phylogenetical
ly
older
than
s
i
r
o
c
e
r
a
i
t
o
p
p
p
n
n
e
i
s
ingeniously
carries
her
prey
underneath
he
r
body
with
p
the smooth dorsal s ide against the earth S he i s s o small
that t he prey must d rag on the ground and i f the ventral
sur face we re against the ground the legs would o ff er re
s i s ta n c e ; so we invar i ably nd her carrying i t w ith the
smooth rounded sur face downward A p roc era carries her
p rey in preci sely the same manner but the point that we
would make he re i s that the wasp i s so large and its legs
s o long that when the caterpillar i s hugged up close to the
body i t i s well o ff the ground s o here w e s ee the uselessness
o f turning the scratchy legs from the ground Yet the
habi t persi sts and w e can see in the habit only the vestige
o f one which was a necess ity in the olde r speci es A p r oc er a
o ften seems only a giant var i ety o f A p ic tip en nis s ince i t i s
p ractically the same i n color form and all respects except
ing si ze One can easily imagine certai n individuals o f the
group
a
l
ittle
large
r
in
s
ize
getting
large
r
cater
t
n
n
i
c
i
i
s
e
p
p
pillars an d the co ns equen t e ff ect o f a su p e rabundance o f
on ,
2 42
food upo n the emerging wasp and its resultant abil ity to
grappl e i n turn w ith large p rey Then when the i n term e
diate f o rms had droppe d o ff there would be le f t only the
two di stinct forms A p i c tip en n i s wh ich uses one medium
s ized caterp illar and A pr o c er a which uses one large one
w ith the i nstincts o f nest bu ilding and carrying p rey not at

all changed
A p r o c era! seems to b e one o f our most husky species for
enduring the col d o f autumn On O ctobe r 2 8 at Cl i fton
Illinois w e took three specimens and s a w many others on the
blu ffs o f th e M i ssi ss ippi The days were sunny but the
nights were very cold
They have many times been seen feeding on the owers o f
white snaker o ot and M elil o tus and the speci es is repo rted
by B anks to frequent the owers o f C ea n o thus The only
enemy mentioned for th is wasp i s the twi sted w ing parasi te
recorded by P i erce
The behavio r o f thi s insect has been worked out by H art
man
H e nds her very tolerant o f human company
wh ich our M issour i wasps are not The work o f excavating
o ccup ies about thi rty v e m inutes ; the gallery i s nearly verti
cal a t the top and runs down i n a gentl e curve until it b e
comes almost horizontal where it w i dens out into a spacious
pocket The large individual s he says store but one i m
mense green tomato c ate rpill a r an d the sm a l ler ones s ev
eral caterp illars H e tel ls how a pai r o f f emal es had a duel
ove r a caterpillar and relate s various pretty incidents
A fte r the c ustom o f many wasps o f h er grou p A p roc era
does not sleep i n i nd ividual nests but assembles i n small
gr o ups to sleep The reason fo r thi s gregariousness in sleep
has not been ascertained We watched one such group o f
these wasps which returned night a fter night to the same
dried stem about th ree feet tall wh ich stoo d i n the m idst o f
abundant green vegetation Th ei r mandibles were l oc ked
.
1 7
1 7
B ull U
.
n i v T ex
.
: 1 1
20
1 90 5
CH A P TE R X I I
S 0M E
Exp er i ments on
S O CI A L W A S P S
he H omi ng o f
P o lis tes p a llip es
The abil i ty o f P olis tes p a llip es to nd its w a y home to the

n est when c a rried i n com plete darknes s and l iberated at
d istant poi nts has proved a fasc i nat i n g and f ru i t ful ex p eri
mental study Fab re found that C erc eri s tub erc ula returns
home i n a fai r p roportion o f i nstances when l iberated two
o r even three k i lometres f rom the nesting s i te H e al so
found that a mason bee exhibits s im ilar abil ity Thes e o f

cours e a re solitary Hymenopte ra It i s indeed m arvel lous
that even a few o f them should return i n cons ideration o f
the fact that these i nsects do thei r foraging nea r home and
i n all p robabil i ty had neve r b e fore v is ited the spo t f rom
whence they were l ibe rated Fabre lays undue stress upon
the fact that s o me o f t hes e insects returned but fails to ex
plain w hy the others d id not do s o H e cannot attribute
the i r return to memory ; but to use his own wo rds : to

some special faculty which w e must content ourselves W i th
recogn iz in g by i ts aston i sh ing e ffects w ithout p re tending to
explain i t s o greatly d oes it transcend our own p sychology

We supp ose howeve r that he i s w ill ing to apply thi s last
statement only to those which made the success ful return
for this explanation vagu e as i t i s doe s not apply to tho s e
which dropped by the ways ide ; fo r th em he o ffers no theory
H e neglects al so to cons ider the age experience and probab ly
the s ex o f h i s subj ects and to make the markings o f i denti
.
S O M E S O CIAL WAS P S
245
cation on the i nsects o f a pe rmanent nature s o that i t

could be certain that any one wasp w a s not accidentally
used i n more than on e ex p eriment
In our work on the homing o f P olis tes p allip es these
factors are al l care fully considered ; w ith permanent mark
ings with consideration o f the age experience and sex o f
the subj ect the re sults that these expe riments show can be
explained wi thout transcending human p sychology
P oli s tes p a lhp es i s a s o ci al w as p having th ree fo rms o f in
d iv id ua ls i n a co lony ; the quee n o r fem ale mal es an d work
ers
They are natural paper makers manu facturing the
paper for thei r nests f rom weathered wood They m ix the
woo dy pulp w ith a ui d substance f rom thei r mouths and
f orm it into the cell s o f thei r nests The nests are made by
a s ingl e colony in one season and all but the fertil ized queen
peri sh with the cold o f autum n At the approach o f w i nter
the fertil i zed females crawl i nto sheltered noo ks and remain
there i n a do rmant c o ndit i on until the warm weathe r o f
spring awaken s them They at once c o nstruct a nest lay an
egg in each cell and when the larvae hatch they feed and
care for them until they trans f o rm into pe r fect i nsects The
rst brood consi sts o f wo rkers only and these at once take
up the task o f enlarging the nest and caring for the larvae
f or a fte r thi s rst brood has developed the queen d o es noth
ing but lay eggs The workers a re undeveloped female s and
i f the que en d ies the bu rden o f egg l a ying i s assumed
by one or more o f the workers Late r in the season the
queens lay eggs wh ich develop into pe r fect males and queens
These mate the males and workers die and the females
s
hibernate
Lewi
)
(
Throughout these experiments i t i s well to remembe r this
important point : that the queens spend the w inter i n some
sheltered nook and hence hav e had some exper i ence not only
i n the previous autumn s roaming but al so in ying about
,
'
2 46
looking for a place o f hibernati on and duri ng the sunny

days o f autumn they O ften come out o f their no o ks fo r play
and sunshine ( s ee page
Furthermore i n sp ring when
n i di c a t i o n begins they y about in search o f p ulp for the
nest insect food for the larvae and nectar fo r themselves
All o f these things are mentione d merely to show ho w i t i s
poss ibl e f or these activities to add to the topograph ical ex
r
e
i
n
e
c
s
h
e
o
f
the
wasp
s
and
o w age i s a factor worthy o f
p
cons ideration in exper i ments o f th is sort A fter hib ern a t
ing the queen s begi n ne st building in Apri l o r early M ay
and work all alone unti l the wo rke rs emerge about the end o f
June During these tw o months i t i s o f l i fe and death s ig
n i ca n c e to the brood that th e queen learn the env i rons o f
her nest
The rst ser i es o f exper i ments were made w ith queens
At thi s time June 1 9 30 the nests were i n an i deal cond i
ti on ; each nest had f rom twelve to s ixteen cells some with
pupae some w ith large larvae some w ith an egg and still
others un lled All o f this i s an index to the queen s a c t ivi
ties outs ide the home
An item o f value too was that w ith o n e queen on each
nest and the nests d i stributed i n th e dozen o u tbuildings
usual to a farm yard no markings we re needed and a se ries
o f markings was u s ed only i n late r experiments a fte r the
workers had em erged
The queens were usually gently
shaken w ithout handl i ng into a w i re y trap and in com
d
rkness
carried
to
a
di
stant
po
int
hence
the
factor
o
f
l
t
a
e
e
;
p
possibl e i nj ury w a s el im i nated
I n al l o f the exper i ments but one the mass o f cages w a s
given w hi le mov ing and sti ll i n c omplet e darkness a series
o f turns i n eve ry conce ivabl e di rection the more to con fuse
the inmates sens e o f di rection o r l oc ation
The work was carr i e d on at a hill top farm twenty miles
south o f St Lou i s A h a l f mi le ea st o f the hous e ran the
,
2 48
i t described a quarte r ci rcle and f
rav ell ed three

leaving here
hundred f eet i n a northw esterly direction where it alighted
i n anot her tree and was lost to sight No 6 went di rectly
northward and landed i n an oak tree ; i n leaving here i t
turned its course abruptly towa rd the southeast and w a s
s oo n lost to s ight
The great variat i on i n the i n itial di rect i ons o f the ir
i ght and the marked d i ff e rence i n the i ntervals at w hich
these queens returned would tend to show that the po int
o f release one eighth m ile f rom the i r home w a s new to the ir
experience and that they reached hom e by v irtue o f the
method o f t rial and e rror or p erseverance
O f cours e i f they ew above the tree top s they coul d have

seen i f they had so great a range o f vi si on the conspicuous
group o f farm buildings wh ich was thei r home ; but at that
height they c ould also hav e seen a strikingly s imilar grou p
o f buildings at th e sam e distance westward wh i ch they
m ight eas i ly have m istaken for the i r o w n B ut regardless
o f the cause or the method i t i s wonder ful to kno w that
every one o f these s ix queens made the return ight in f rom
2 2 to 72 m inutes
Thi s ex p e r i ment shows at least that i f
some unknown force draws them back to the nest th i s force
does not act by the m o st economical method as to e ither
time o r space
,
E xp e ri m en t I I
June 2 0 Th i s experiment w a s made w i th n i ne wasp s

wh ich were carried to B ecks The distance along the road
was about 2 % m iles but the distance as a bi rd i es was
app roximately 1 % miles B oth the home and the point o f
rel eas e were o n a b out the same level ; thi s elim inated the
factor o f v a r i ed el ev a t i ons F i ve o f the w a s ps nos 2 3
and
had
been
used
i
n
experiment
I
where
they
had
6
4 5
been em i nently success ful i n mak i ng the one eighth mi l e
.
2 49
ight ; the other f our wasp s 7 8 1 0 and 1 2 had never

be fore been used
These n ine wasps were gathered between 8 :5 0 and 9 :2 5
a m and placed i n indiv i dual large test tubes plugged w ith
l oose cotton the tubes placed unde r cover and carried along
the road to B ecks The contor tions o f the w inding road
we re even i ncreased by the usual turn s o f con fusion All
we re released at one place and time between 1 0 :5 0 and
a m The tempe rature w a s about 9 2 F with no
perceptible w ind In order to reach home they would have
to y southeast
The return o f the ve wasps wh ich had ha d p revious
expe ri ence in the rst trial w a s a s follow s : no 2 when re
leased ew north northwest and al ighted on a telegraph
pol e then it made a large ci rcle and dashed away southeast
toward home but it never arrived at its nest dur i ng the eight
days we kept watch for i t No 3 ew southeast for fteen
feet and alighted on a bush then ci rcled about and ew away
north nor thwest i n quite the o pposite d i rection from he r
home S he returned to the nest at 4 :0 5 p m having made
the trip in ve hours and being the rst to reach the nest in
this experiment as well as t he p receding one N o 4 ew
s o uth southeast to a bush fteen feet away rested for a few
seconds and then ew d i rectly eastward almost the proper
di recti on for reaching hom e Thi s on e di d not arrive at the
nest unti l
the next mornin g a fter an absence f rom
home o f 2 3 hours No 5 when l iberated ew for twelve feet
south southeast and alighted for a m oment on a bush then
turned its course northeastward This one did not return
that day but the next day when our pat i ence w as almost ex
p m a fter an absence o f
ha us ted s he came in at
approximately one and one hal f days N O 6 ew di rectly
westward to a bush twenty feet away Th is one neve r
returned although i t too was sought for eight days
-
25o
Thus o f the ve queens that had made tlfe one eighth

m ile t r i p three success fully made the one and one hal f m ile
ight What these three d id the other two should have done
also s ince thei r d ome stic interests were alike ; each ha d f rom
a twelv e to a s ixteen celled n e s t at hom e to care fo r which
really indicates that all h a d had abo ut equal ex p e rienc e
abroad For an explanation o f the n on appearance o f these
tw o w e can only susp ect that they fa iled in a case i n trial
and error The f acts that n o 2 made the rst ight speedily
that s he l e ft the tube apparently i n good health and vigor
and that by reason o f the aggre ssive coloration and severe
sting enemi es are few would all indicate that her non
return was probably due to he r s imply having gotten lost
I n the cas e o f no 6 how eve r one m ight j ustly suspect that
he r failure to return was due to her subnormal ity since i f
s he had real ly been normal
s he could p robably have made
the ight i n Ex p e r i ment I i n the same time as the others i n
stead O f thrice the time Whether o r not the three which
returned di d s o by reason o f thei r previous experience w ill
be i ndicated by the behav i or o f queens no 7 8 1 0 and 1 2
none o f which had been used i n ights previously
N o 7 when released at 1 1 02 alighted on a bush fteen
feet aw ay an d the n ew aw ay to the southeast I t returned
to its nest the next day at 9 a m a fte r an absence o f a l
most one day
No 8 ew north northwest and al ight ed on a bush fty
feet away I t al so reappeared at the nest the next day at
having
made
the
j
ou
rney
in
p
ractically
the
same
time
:
9 45
as the p rev ious on e
No 1 0 when l iberated alighted on the nge r and remained
there for fteen m inutes when it was placed on a ower
head Thi s one never returned to its nest but its lan guo r
indicated an il lness
N o 1 2 when released s at on the neck o f the b ottle for
-
252
cause on one s ide were heavily wooded hilfs and on the

other the bottom lands between the track and the r iver
were cultivated elds and forest
The wasp s were released on the rail road track j ust at the
edge o f the town
N o I ew north thirty feet made a c i rcl e to the no rtheast
and landed high i n a cotton w o od tree Th i s one never re
turned to the nest
No 3 w a s released at 1 0 :2 5 ; i t ew north northwest and
settled in the weeds at the foo t o f the rail road embankment
I t returned to the nest th e next day at 2 :30 p m
No I I di s a ppeared down the steep embankment as did
no 3 but never returned to the nest
N o I 3 c i rcl ed above our head s ew so uthwest and alight ed
i n a thicket o f sw eet clover
Thi s one neve r returned
to the nest
N o 1 6 beh a ved i n the s a me m anner as 1 3 excep t that
it made its to ilet fo r four m inutes then ew northwa rd and
was lost to sight It l ikew i se did not return
N o 1 5 rst ew sou thward for a few feet then curved
to a southweste rly course and was lost i n the thicket o f
weeds at the foot o f the embankment Thi s one reappeared
at the nest the same day at 7 :30 p m a fter only n ine hour s
on the lon g j ourney
O f the s ix wasps wh ich were started i n thi s tw o m i le race
tw o n ished n o 3 th e o n e wh ich had been o ut on two
p re vi ous test ights and no I 5 out fo r the rst time I t i s
i nteresting to note further that the novice made the i ght in
the shorter time nine hours whereas the other on this her
thi rd ight took more than a day for i t O f course the
factor o f fa tigue may ente r i nto the case o f n o 3 C o ncern
ing the four whic h failed utterly w e can only susp ect that
1
they tr i ed but lost thei r l a ndm arks
-
we
s i s ta n c e
a re
i n d eb t ed to
i n the
e a rly
C a rl
p a rt
of
Ec
a rt
thi s
an d
w k
or
C a rl Wi tter
f or
va luab le
as
253
E xp eri m en t I V
June 2 5 The results o f thi s experiment w i ll have to be

ignored but are recorded here to save con fusion later In
thi s test queens 8 4 and 1 5 were l iberated from a moving
train
S ince they never c ame home w e do not kn o w
whethe r they died or got lost as did the others o r became
entangled in the whirlw ind and w ere d ra wn unde r the
wheels Neve rtheless l ike the noveli st w ho must grace fully
di s p ose o f h is characters when they are o f n o further use so
let this record di spose o f queens 4 8 and 1 5 The nests
wh ich they le ft beh ind were o f course therea fter orphan
nests ; we shall have more to say later o f the orphan workers
.
H O M I N G OF W OR K E R S AN D
THE
EEN
S
Q
U
The workers we re beg i nning to emerge Thi s n ec es s i

ta ted the adding o f marks o f identi cati o n on thei r bodies
Thi s w a s ve ry art fully done without inj ury to the insects
Fabre attributes the fai lu re o f m any o f hi s i nsects to return
to the nest to inj ury i n handling when being m arked B y
our method inj ury was avoided and i f an ind ividual failed
to return the re was n o cause to bl a me but its own stupidity
o r its youth fulness In taking the wasps fo r marking a w ire
y trap was usually placed over the nest and the i nmate s
shaken into it carried thus t o the laboratory permitted to
esca p e one at a time to a w indow pane where they were
p icked up w ith a wad o f so ft cotton and gently held b e
tween the ngers whi le paint was a p p lied i n the des i red
colors and numbe r o f dots The markings on the thorax
were most practical fo r here they were reta i ned for l i fe
whereas on the abdom en i t was possible for the wasp s to
clean them o ff a fte r several days
.
254
E xp eri m en t
June 2 5 The queen o f nest 2 failed to return to her nest

i n Experiment I I on June 2 0 The pupae were able to con
t in ue thei r development w ithout he r howeve r and on June
2 4 two worker s eme rged and one on Jun e 2
5
O f course i f these wasp s nd their way home i n st in c
t iv ely o r through the aid o f a s ixth sense w e have a right to
ex pect the young wasps t o return as readily as the ol d ones ;
i f howeve r age and experience are factors i n di recting
them home the edgl in g s woul d be unable to return to the
nest s ince they lack both
These three we re placed in large test tubes and w ithout
the hand icap o f darkness s o they coul d have every advantage
o f studying the ir landmarks they were taken to the mail
box one eighth m il e di stant along a level road ( same as
Experiment I ) an d libe rated a t
a m They le ft the
tubes rather awkwardly ( the di rect activity and impatience
fo r e s cape s o n oticeabl e i n th e queens wa s absent ) and
settled on the vegetatio n below
Fou r days o f care ful watch ing at th e nests showed the
continuous absence o f these workers They were all un
abl e to nd thei r way home desp ite the short di stance and
the f ree use o f the i r sense o f sight
.
VI
E xp eri m en t
July 7 N O further ex p e rimental work was done on the

wasps b etween the last test on June 2 5 and the present one
on July 7 I n the i nterval several workers had emerged
f rom the various nests Thi s experiment i s th e same as
Experiment V except that a large r number o f wasps w a s
used wh ich ought to give more conclusive evidence as to
whether the hom ing process i s one o f learn ing o r o f in
Thus ten w o rkers and
s ti n c t o r o f some unknown sense
.
256
erect ; at a nearer app roach they y downward about the

head o f the i ntruder and o ften are f ree w i th stings A fte r
a few seconds o f aggress ive buzz ing most o f them escape
to return later ; those wh ich remain are usual ly the queen
and the youngest w ork ers I t seem s that the young work
ers are sl o w to learn to de fend themselves by o ffens ive
actions True they can sting i f handled but they do not
y away f rom the nest when it i s i n danger nor do they
attack an observer but remai n on the nest When teased
with a straw they walk a short distance and stop ; when
followed they escap e by the rst crack i n the wall I n our
bservat
i
ons
these
n
wly
hatched on es when o usted i n thi s
O
e
manne r seldom were able to return to the nest this fai l
ure to return ev i dently b e i ng due to the i r lack o f knowl edge
o f the i r env i ronment
N o w th i s w a s prec i sely the condit i on i n nest 1 9 in
Experiment VI ; the ol d workers ew at our approach and
the queen and three young workers rema i ned and we re
taken for the experiment The queen returned and these
new wo rkers w ere all lost ( S ee Expe riment VI I for addi
t i on a l observat i on s o n workers o f nest
.
E xp eri m en t
VI I
Th i s i s merely a repetition o f the fo rego i ng ex

h
A
second
lot
o
f
wasp
s
we
re
l
iberated
at
t
e
ri m en t
e
p
sam e place and date a s be fo re but two hours later at
1 0 :30 a m
Nest 1 9 A fter the rst lot had been taken and released
the O lde r workers which escaped us r et urn ed to the nest
Three o f these were taken and marked di ff erently to d is
t i n g ui s h them w ith ce rta i nty from th ose f rom thi s nest us ed
i n p revious exper i ments and released as per account
When the nests were exam i ned at
O clock two o f
these workers had al ready returned wh i ch sat i s es us i n th i s
July 7
57
case as in others to follow that the w orke r wasp wh ich i s

O ld and experienced enough to escape or show bel licosity
when taken has by that time the abil ity to return to the
nest
Nest 1 3 This queen w a s lost on June 2 1 in Experiment
I I I and on thi s date som e hal f dozen orphans had charge
o f the nest S o ale rt were they that only one o f them could
be taken It never returned to the nest
Nest I 5 From thi s nest we have some intere sting data
The queen was l o st i n Ex p e riment I V on June 2 5 The
queenless nest w ith its potential l i fe was taken from the barn
on June 2 7 and p inned to the wall in the labo ratory near
the tabl e where we could watch the maneuvers o f the young
when they emerged I t also happened that on o r about
June 2 2 we had been experimenting on wasps in applying
paint to various parts o f the body and observing its e ffect
upon the in sects On that day two workers p robably a day
or two old were taken from nest 1 2 in the barn carried into
the laborato ry di rectly i n f ront o f the barn at a di stance o f
fty feet The re two d o ts o f paint we re appl ied to the
pro thorax o f each and a fter noting the meth o d o f handling
thei r abil ity to clean o ff the paint etc they were l iberated
seven hours late r by being allowed to walk ab o ut On the
S ill o f the open window nearest the barn We expe cted that
o f course they w oul d nd thei r way back to the nest in
the barn di rectly in f ront o f the window and only fty f eet
distant but somehow these workers did not nd nest 1 2
We l ittl e suspected that thi s new envi ronment as they
could s ee i t through the w i re y cage during the seven hou rs
o f impri sonment and thei r subsequent freedom about the
room was making such an impression upo n thei r senses that
they would regard i t as thei r hab itat rather than the barn
Five days later orphan nests 8 and 1 5 were p inned to the
wal l near the laboratory table where these worke rs had
,
258
been handled Then i t became evident that th ey must have

been l inge ring about th is building while w e l oo ked else
whe re fo r them for on the same day that these two nests
were p inned to the wal l s ide by s ide both o f thes e e s pe
c i a l ly marked wasps we re found at rest on nest 8
where
they remained p ersi stently for three days when one shi fted
to nest I 5 and each hel d its res p ective nest continu o usly
therea fte r Th is sel f appointed queen on nest 1 5 took care
o f the b r o od depos ited eggs fed the larvae tore down parts
o f the nest and rebuilt other sections O n June 2 9 one
worke r eme rged and on the following day another When
all three o f thes e we re used in thi s experiment on July 8
the worke r m o the r w as at lea st seventeen days old and the
wo rke rs ten and n ine days ol d respectively Ow ing to the
sm allnes s o f the col ony they must have had w ider out o f
doo r expe riences than the workers o f large r colonies
H ere i t i s we ll to m ention in anticipation o f what we
shall s ee i n later experiments that age and expe rience shoul d
not be taken as synonymous
We think o f course that
old workers woul d be more ex p e ri enced than young ones
b ut thi s i s true only up to a c ertain point
When the colony
i s growing a supply o f pulp fo r the nest and food for the
many young mouth s makes i ntensive wo rk fo r the small
numbe r o f workers A fte r the season has pas s ed its height
or the young are few i n numbe r with an abundance o f
workers they may spend more and more time on the nest
and when occas ionally they y abroad they remain in the
general vicinity o f the nest seeking l iquid re freshment F i
nally in the early autumn one sees the cells enti rely devoid
o f young and wo rke rs mal es and late r f em ales clinging
languidly to the carcass o f a nest
B ut now to resum e the n a rrative o f Ex p e riment VI I the
three p ioneers on nest 1 5 were rel eased with the othe rs at
the mail box at 1 0 :30 o clock All three o f these returned
.
2 6o
VI I I
E xp eri m en t
July 9 This test i n all ess entials the sam e as Experiment

VI I was p e r fo rmed w ith v e workers f rom two o rphan
nests th ree workers from n est 1 8 which emerged Jun e 2 9 to
July I and two f rom nest 1 1 which also emerged June 2 9
an d July 1 These were tak en to the mai l box and released
at 1 0 :2 5
Two o f the workers from nest 1 8 returned at 1 1 I o and
and two f r o m nest 1 1 one at I I 5 5 a m and the
othe r at 1 :35 p m H ere we s ee worke rs returning w ith
eminent succes s
We shall note two things i n passing :
they we re all e ight to eleven days old and they were sol ely
respons ible for the maintenance o f the home and hence
had probably had wider experi ence i n th e wor ld tha n i ndo
lent youngsters
The reader wi ll rem embe r that young
workers read ily lose themselves even though l iberated near
the nest
Up to the p resent point in o ur s tudi es ol d workers have
success fully made the tri p ove r on e e ighth m ile di stance
and queens have return ed from a two m i le tri p but the old
worke rs hav e not yet been experimentally tested in long
ights H ence th e fol low ing experimen t was c arried on
w ith the purpo se o f trying workers both old and young i n
a ight o f longe r distance
.
E xp eri m en t I X
July 1 7 The wasps were taken f rom the buildings about

the farm and two l ots nest 2 4 and 2 5 were brought from
a shed near the railroad track and carried up a h ill o f per
haps I 5 0 feet el evation to the laboratory to be marked
Thi s was done at noon Later they were taken down the
gently sl o p i ng h ill side by an entirely d i ff e rent route to the
rail road track and then carried to M ontesano P a rk on the
.
26 1
outski rts o f Kimmswick The wasp s all experienced the

same conditions except that those from nests 2 4 and 2 5 had
in addition to the trip to Kimmswick a trip by a tortuous
route up the hill to the laboratory At 3 :4 5 p m twenty
two wasps were l iberated at M ontesano P ark one and eight
tenths miles south o f thei r homes and 1 5 0 feet lower for
all but those f rom nests 2 4 and 2 5 The cages were dark
ened and turned about repeatedly the more to con fuse the
inmates
Three wasp s from nest 2 5 were taken Th is nest was
large having ten larvae ready to spin thei r coc o ons s ix
w ith sealed cocoons and numerous young
One would
suspect that by the time the proprietors had accompl i shed
thi s much work they would have had su fcient foraging
experience to be able to get to thei r homes f rom a point
one and eight tenths m ile s di stant on the same level Only
one o f them returned ; it appe ared on the next day at 2 p m
We suspected that thi s one was the queen but had no
proo f o f the fact
The ages o f the other two were
unknown
Nest 2 4 w a s near nest 2 5 i n the shanty by the railroad
track Five wasps we re taken fr om this each marked w ith
two white dots on the p ro tho rax O f the se only one re
"
turned ; i t appeared the next day at
p n1
Nest I 5 The worke rs from thi s nest were the th ree
wasp s which had taken over the queenly duties and had
succes s fully made the one eighth mil e trip in Experiment

VI I This had been about ten days previously ; the addi
t io n a l time for becoming acquainted w ith the neighborh o od
should have been a factor to aid them in making a p rompt
.
S i n c e d uri n g l a te r
n es t s ,
w k
or
th e s e t w o
w p w
y w
qu
w e ha ve f ul ly d ec i d e d t ha t t h e
p e r i m en t s
re
f er
to t h e m
as
q ueen s
as
e re
c o n s t a n t ly
e re s o
e en s
an d
on
i n l a te r
th e
ex
2 62
return H owever none returned from thi s greater di s

tance
N est 1 9 gave us two worke rs o f unknown age but each
bearing a white mark i ndicating that it had made the one
eighth m ile ight previously Ne ithe r o f them came back
Nest 1 1 The one wasp which was taken f rom thi s nest
still bore the m a rk s w hich i denti ed it as the one used i n the
mail box trial on July 9 ( Exp V I I I ) Desp ite its prev iou s
success ful trip i n Expe riment VI I I it did not return to the
nest this time
Nest 1 8 Just as in the p receding i nstance on e w orker
which had success fully made the trip i n Expe riment V II I
was again taken but thi s time i t never reappea red at the
nest
N est 2 2 w a s a newly di scovered nest in the smoke house
which had only one adult tw o s ealed c ells and a hal f dozen
other young Th i s sole p rop r i etor p resumably the queen
was taken on this o urn ey but never returned
Nest 2 0 A young worker ve days old was taken on
th i s long trip but as w a s to be expected it d id not reappear
From nest 6 thre e wasp s whose markings p roved that
t hey had success fully mad e the one eighth mile trip on July
8 ( Exp VI I ) w ere agai n used in thi s expe riment but n o t
one returned th i s time
Nest 1 3 Tw o workers o f unknown age were both lost
I n thi s ex p erim ent i t i s at o nce evident that no worke r
taken out for the rst time was able to nd its w ay home
ove r a di stance o f one and e ight tenths miles and even other
workers and worker queens although they had success ful ly
made other return ights we re unabl e to do s o at thi s
di stance
It might be suggested that the meteorological
conditions on th a t day we re p e cul iar in some way hindering
thei r activity but we must not los e sight o f the fact that
the queens f rom nests 2 4 and 2 5 actin g as a check upon
.
2 64
Xb
The wasps from nests 2 4 and 2 5 for Ex p e riment I X had
been removed on the preceding day ; they were all still absent
from the nest at 7 o clock on the morn ing o f July 1 8 B ut
those wh ich had escaped the rst c ollection had by th i s time
returned and four o f these were taken The very fact that
these escaped from the n est on the p revious day when the
catch w a s made indicates that they were the older and w ise r
heads and the results should be regarded in thi s l i ght One
These
w a s taken f rom nest 2 4 and three f rom nest 2 5
were rel eased on the h i ll top at 8 o clock and when w e ar
rived a t the shed below at 8 :5 0 they had all retu rned more
speedily than w e
I t i s too great a deduction f rom l imited data to s a y that
i t i s eas ier for the wasp s whose home i s i n the valley to
reach it f rom a h i ll top than v ice versa but the experiment
does show that i t is possible fo r wasps to nd the i r w ay
home f rom hill top s i f they are taken from thei r homes
below regardless o f what b ees do I n Experiment IX where
they returned f rom a l ower plane to the sam e hill top the
route was by a two m ile ight and the ascent w a s a grad
ual one up the gently S lop ing s ide o f the hill whereas in
thi s e xperiment the ascent would have been up the steep est
si de o f the h ill
E xp eri m en t
E xp eri m en t XI
Wh ile we were eager to make some ex p erim en ts o n the

homing ight o f these wasps from greater d istances these
had to be postponed as the material avail able at thi s time
w a s unsuitable
I t seemed ttin g the re fore to devote the avail able material
to experiments attempting to ascer ta in by what sense they
'
ma
III
E p er i m en t
in
th e
ow w ar
k g
s ho
ig ht
t h at
a n t e n n ae l e s s
s uc c e s s ul ly
a sp s
a re
c a p a b le
of
265
nd the i r way hom e It was our intention to depr ive them

o f the use o f the ir eyes or antennae and then test th eir abil
i ty to make return ights which normal individuals had
p roved themselves capable o f making The antennae we re
removed in part o r whole one o r both To take such a d
vantage o f the fascinating creatures which seem alm o st pets
is degrading indeed ; one could d o no more and the further
trick o f blinding the eyes o f such highly endowed beings
w a s abandoned
.
E xp eri m en t XI a
Jul y 2 3 The obj ect o f this experiment was to test the

abil ity o f a n ten n a el es s wasps as compared w ith normal
indivi duals to make the h o ming ight For thi s purpose
we desi red a random sampl e o f the populati on from whi c h
all the naturally unt individual s were el iminated This
end w e accomplishe d by merely sending a lot o f marked
wasps on the ight f rom the mail box and then taking those
which were success ful in this one eighth mi le ight fo r fur
the r experimentation I t seemed a poo r reward indeed for
thei r faith fulness and me ri t merely to be permitted again

to make the involuntary j ourney plus mutilations or rather
m inus parts o f the antennae
Al l o f these unmutilated wasps were taken between 8 and
o
clock
marked
carri
ed
to
th
e
mai
l
b
o
x
in
th
e
usual
man
9
ner and liberated at 9 :4 5 a m
Three workers we re taken from nest 1 0 The queen from
thi s colony had been lost on June 2 0 but the workers had
done wonder fully well and unde r thei r care the nest had
reached very ne pr o portions Under thes e c i rc um s tan c es
i t i s not surpri sing that these three wo rkers had returned to
the nest by the time w e arrived to examine it at I I 3o
Nest 2 6 contributed three worke rs and a queen We
suspect that the workers may have been young because the
.
'
WA S P S TUDI E S AFI EL D
2 66
older and mo re expe rienced ones w ere a g ile i n escapi ng

when we approached to m ake a c atc h At 1 1 30 only on e
m
the
qu
e
en
had
returned
The
next
day
at
p
(
)
whil e waiting fo r the mail w e s a w one o f the marked
workers sitting o n a p ost fty feet f rom the p lac e wh ere
i t had been l ibe r a t ed W e thou ght at rst that i t w a s i ll
or inj ured but when app roached it darted n imbly away i n
apparent goo d health I t may have been young o r too i n
d i ffe rent even to start back home I n any event it never
reached home
The one worker two days old f rom nest I 3 did not return
The queen f ro m nest 2 0 returned at
p m
E x p eri m en t
XI b
We were now ready fo r the second step i n the ex p eri

ment For thi s purpose the above mentioned success ful
comp etitors and a few others which had p roved capable
i n p revi o us ights together w ith a small number o f en
t i rely new and inexpe rienced wasp s all d istinctly marked
a n d w ith vari ou sly modied antennae
we re liberated as
usual at the m ai l box on t he sa me a fternoon at
Thi s allowed them ampl e tim e to return be fore night fall
thre e hours late r When these wasp s were released they all
ew away w ith the usual alertness and energy s o one s ee
ing them would susp ect that the lo ss o f thei r antennae made
no di fference to them
From nest 1 8 two workers were taken and the le ft anten
na removed near the b as e
One o f them bore a white
ma rk p roving that it had been success ful in a previou s
ight ; the othe r was new i n the work The next day b e
tween 7 :30 and 8 :30 b oth o f these ha d retur n ed
T w o workers f rom nest 1 1 w ith p revi o us experience
were each dep rived o f the right antenna and sent out on
-
La t e r
o b s e rv ati o n s
p r o v e d t hi s
t o b e t h e q ueen
'
o ne
2 68
w ith
both antennae amputated at the base and anothe r

wh ich had been success ful in returning w ith on e antenna
in Experiment X I ; its othe r antenna w a s n o w cut o ff The
one us ed fo r the rs t time d id n o t return to the nest but the
other n ow repeated the trip dep rived o f both these organs
She did not a rrive at the nest however until 1 00 p m
Nest 2 0 gave us one queen which had success fully made
this di stance i n Exper iment X I She was now handicapped
by having on e hal f o f each antenna cut o ff and l iberated
w i th the others for the same ight S he neve r returned
From nest 1 0 two wasps were taken wh ich had p re
v i ously made the one eighth mi le ight success fully when i n

a normal condition Now they we re l iberated agai n w ith
one hal f o f each antenna rem oved B oth o f them returned
between 1 1 and
a m A third worker f rom th i s
nest a edgl ing also w ith one hal f o f each antenna re
moved did not return
From nest 1 9 a worker which had been success ful in
on e o f the previ ous tests was t ri ed i n thi s ex p e rime n t w ith
its entire right antenn a gone This one returned to the
nest at
a m
The i nsects in the forego ing tests were all l iberated at
:
m
a
those
wh
i
ch
follow
we
re
released
on
the
same
0
;
7 3
day and place at 3 : 5 0 p m
The two wasps f rom nest 1 1 used i n Experiment X I
where the right antenna w a s removed we re now sent out
w ith both organs gone B oth returned to the ir nest during
the early part o f the a fternoon o f the follow i ng day
The wo rke r f rom nest 1 9 wh ich success fully returned
minu s the right antenna in the rst part o f thi s experiment
w a s now again tried i n the same ight w i th th e othe r t e
moved I t reap p eared at the nest seem ing very sp rigli tl v
and unhampered in only forty m inutes !
T he same t reatment was now accorded to the two wasps
,
'
2 69
S O M E S O CIAL WA S P S
f rom nest 1 0 which w ith one antenna had been success ful
in Experiment XI When the nest was examined at 4 :30
one o f these had already returned ; the other did not arrive
h o me
To summarize then : the wasps in thi s lot which had been
p reviously success ful i n the ight w ith only one antenna
now p roved themselves equally capable w ith both antennae
amputated These were subj ected to all the con fusion that
the o thers endured i e the darkness and the turning
about
The time i n which they m a de the ight was a
l ittle longer than the time required by the rst lot o f queens
in Experiment I ; the m inimum tim e for these w o rkers w a s
o
0
minutes
and
two
we
re
out
ab
ut
2
0
hours
4
We had planned to experiment upon these wasps entirely
o r partly a n ten n a l es s w ith ights o f greate r di stance but
were called away from July 2 5 to 30 We thought furth er
more that a rest fo r these overworked ind ividuals would
el iminate the factor o f fatigue f rom the results o f long
ight experiments We assumed that si nce most o f them
returned very p roperly to the nest even under hand icap they
w o ul d o f cou rse continue the i r normal activities until our
return B ut at the end o f the week when the various nest s
we re examined not one o f either the partly o r entirely mu
t ila te d wasps w a s to be found
Whether they had per
i s hed or had been driven from the nests by the normal
inhabitants or had lost themselves in foraging expediti o ns
we do not know Thus end e d the plans f o r thei r long d i s
tance ights under handicap
,
E xp eri m en t
XI I I
July 30 I n Experiment X the wasps in no rmal condi
ti o n were taken from thei r outbuilding at the foot o f the
hill and carried to the crest o f i t whence they all speedily
.
2 7o
made thei r way home The expe riment was h o w repeated

with wasps which had the antennae cut o ff at the base
Fou r wasps we re taken f rom nests 2 4 and 2 5 and released
at I 2 o c l o c k Two were unmarked show ing that they had
not been used previously ; a yellow dot wa s placed on the
thora x o f each Two others al ready bore the white mark
wh i ch showed t hem to h ave had s ome previ ou s acquaint
ance w ith thei r tormentors Two o f the f our wasps returned
to the nest ; one w ith a white mark reappeared at 4 p m
and the other a novice returned home at 5 :3o
.
V
S eptember 3 Du ring the preceding week mal es had been
appear i ng on the nests T hes e w ith what wo rker s were
available compo sed the material for th i s expe riment S ince
the males had no i nte rest i n the i r nests one woul d only
expect them to get lost under such treatment A fter the
males have emerged from the nest they are p ractically done
w ith it s ince the i r function is merely the ferti li zation o f
the femal es and fo r thi s purpose they undoubtedly go else
whe re and do not need to remember the hom e nest
The wasps were captured at 2 p m carri ed to the labora
to ry and marked but on account o f rain the ir release at
the mail box was p ostponed until 4 o clock A fter that hour
i t was cloudy and dreary and much rain fell during the
n ight ; the next morn ing also w a s cloudy but in the a fter
noon the s un shone out b rightly
o
f
Nest 2 7 was out
doors on a rose bush at the foot o f
the hill Thi s gave us one queen and on e worker all used
for the rst time These we re taken to the top o f the hill
again marked a nd rel eased B oth w o rke r and queen were
found to have returned the next day at 9 : 1 5 a m
N est 2 8 w a s under the porch bes ide 2 7 Two workers
were taken f rom thi s and marked yellow One had returned
E xp eri m en t XI
2 72
i n Experi ment X and one worker w ith a yellow mark

used i n Experiment X I I I on July 30 At 1 1 a m the
white marked queen returned ; at 2 p m the one w ith the
yellow mark appeared and at 2 :30 anothe r worker which
had neve r be fore made the test ight returned
N est 2 7 one worker used for the rst time returned
at I I 1 0 a m
N est 2 4 three workers new to these experiments o f
which one returne d at 2 p m
N est 2 8 one male and four wo rkers used for the rst
time ; none o f these r eturned
,
E xp eri m en t X VI
S eptember 5 This was an experiment in long di stance

ight w ith whatever material th e lateness o f the season
a fforded The wasps were gathered between 4 and
r
m
taken
to
the
labo
ato
ry
on
the
h
ill
top
S
eptember
4
p
marked fed on honey o f which they ate greedi ly kept
i n w i re cages over n ight and l iberated the next day The
mater i al from nests 2 9 to 33 was taken f rom an ol d shed
n ear the rail road track one fourth mile south o f the labora
tory ; thes e had not been used exp erimentally be fo re On
the morn ing o f S eptember 5 under p rop er wrapp ings they
were al l taken on board the 7 :2 5 train to a station
m iles
to the north removed from the train and released at 8
The s un w a s shining warm and bright and a
o clock
.
T h e m a teri a l w a s
w hit
e
u sed
e,
x t ra
XV
y e ll o
an d
c e s s f u l ly
us ed
f u l ly
26 :
on
in
s uc c e s s
f ul ly
d ot ; N e s t
in
Exp
f o llo w s : N e s t
in
E xp
w k
27 : 2
or
X I IV ; N e s t
XV
e rs ,
23 :
an d
I u
w k
or
se
k
w
O n e q ue e n m a r e d i n
n o w m a r ed
ith a n
25 :
e rs ,
s u c c e s s u l ly
l
l
e
o
y
in
us e
Ex p
s uc
w k
i n E x p X I V, a n d 2 n ev e r b e o re u se d ; N e s t 2 4 : 2
o r ers ,
r er
1
o
us e
l
x
X
e
:
1
s uc c e s
s
r
i
E
N
t
e
v
o
u
s
i
n
V
;
p
9
p
y
E x p XI V ; N e s t 2 : 3
o r er s n o w u s e d f o r
rs t t im e ; N e s t
w k
w k
I w k
or
e r,
a b d o m en ,
t o 33, 9
as
w k
or
not
u s ed
b e f o re
u s e d s uc c es s ul ly
e rs
an
m al e
an d
in E x p
e re
u se d,
XI
all
w i th d t
th
J u ly 3 ; f m
f
th t t i m
q u een
on
or
on
o ra x
n es t s
ro
rs
an d
29
2 73
breeze s o gentle as to be almost i mpe rceptible came f rom

the south All o f the wasps were from places at the railroad
level the same elevation as the po i nt o f release exceptin g
those f rom nests 2 1 9 and 2 6 ; these were hill top nests at
an altitude o f I 5 0 feet highe r The Meramec Rive r inte r
s ec ted thei r course o f ight nea r home
These twenty four wasps many o f which had records o f
succes s ful ights ( s ee f o otnote page 2 7 2 ) we re liberated at
8 a 111
O n account o f the di stance w e expected that none
would return ; hence we were almost shoc ked to nd that
one f rom nest 2 6 had come back I t w a s the queen wh i ch
had nished in the preliminary ight i n Experiment X I A
di stance o f
m iles plus an elevation o f 1 5 0 feet is indeed
an astonishing feat for so small a creature At 6 : I O that
evening one queen appeared on nest 2 5 Thi s wasp had
al ready made two success ful ights Experiments X and
XV Subsequently the nests were watched for a week but
o ut o f the abundance used in thi s experiment only these two
ever returned I t seems that the oldest and most experienced
ones were the most expert in nding the i r w a y home
or the most success ful unde r adve rse ci rcumsta n ces
If
magic o r some impell ing powe r drew the wasps to their
nests as B ethe th inks o f the bees then surely a di fferent
type o f behavi o r woul d be mani fested
,
S U M M AR ! AN D C O N C L U S I O N S
T o summarize then w e nd ( table A ) that out o f 33
queens which were taken fo r various distances from one
eighth m ile to two and seven tenths miles 2 4 returned
to the nest
O f the 2 2 taken out for the rst time 1 7 reached home ; o f
the I I which had made p rev i ous test ights 7 success fully
returned
,
2 74
T A BLE A
H om i ng
q u ee n s
23
'
=3
:3
0
0
1
" C ae
o
:4
n
2
U}
t a
T o ta l s
*
To
v a lley
b elo
W ith the workers the results are quite d i ff erent

O f 1 1 2 workers that we re used on ly 2 8 returned and
tho se wh ich returned d i d s o o nly i n the short ights The
long distance tests always gave negat i ve results Let us
go into detail in regard to these 1 1 2 workers and the 2 8
that returne d home i n thi s lot O ut o f the I 7 new workers
that were experimented upon n one returned ; o f the 33 old
w o rkers 1 4 came back ; o f the 6 2 wo rkers o f unknown age
1 4 returned
A c cordin g to the law o f chance it i s easily
possible that these 1 4 out o f 6 2 worke rs o f unknown age
may have been ol d ones The last four colu m
ns s how that
.
'
2 76
previ ou s exp e rience i s not necessary to succ ess ful ights

There we s ee that 2 7 success ful returns out o f 9 6
were made by wasp s used for the rst time and that
only I out o f 1 6 wasp s that had had p reviou s ights made
a success ful second trip
W ith the q ueens thi s i s not
the case
Tabl e C shows that not one o f the 1 7 males returne d t o
thei r forme r nests even though the distance fo r 1 6 o f them
was only one eighth m ile
Tabl e D sh ows that the function o f the antennae i s n ot
the sole factor in bringing them home for out o f 2 4 muti
lated wasp s 1 8 success fully returned
Thus by th e el im ination o f other faculties the ev i dence
grows stronge r that v ision i s the sense whereby the insects
regain thei r homes
.
TABLE
H o mi n g
f m a l es
a?
"
s
q)
I?
L}
L
a)
f:
XIV
XV
ir
X VI
it s,
38
:
m
3
a
5
ES
!
1
3
1
! I G
T o ta ls
The evidence here shows that P ol is tes p allip es nd thei r

w a y home i n the m anner which Lubbock describes for hi s
bees : I f a h ive bee i s taken to a di stance s he behaves as

a p igeon does unde r s imilar circumstances ; that i s to s ay
s he ies round
and round gradually ris ing higher and
8
S en s es
I n s ti n c t s
an
I n tel li g en c e
of
An i m a l s
26 5
1 888
T A BLE D
H om ing
2 77
f m u ti la ted
wasp s
'
4)
"
'
'
3 33
x
II l
a)
13
3
O
ve
'
d:
0
T o ta l
XIII
90 5
d
D
3%
:
6
8
of
One
t he s e w a s
q ueen
highe r and enlarging her ci rcle until we suppo se her

stren gt h fai ls or she come s w ithin sight o f some known
obj ect
Ag ain i f the bees had returned b y a sense o f di r ection

they would have b e en back in a few minutes To y one
and a hal f o r two miles would not tak e ve minutes One
bee o ut o f the 1 4 7 di d it i n that time ; but the others took
one two t hre e or even ve hours Surely then it i s rea
sona b le to suppose that these lost some time be fore they
came i n sight o f any obj ect known to them

Fabre has done some work on the C er c eris wasp and on
two species o f Os m i a bees insects which while l iving in

communiti es are sol ita ry in habit In al l three species a
cer tain propo rtion o f those experimented upon returned
to the nest a fter having been l iberated at di stances varying
from two to three miles fr o m thei r homes He cannot
comprehend the p o ssibil ity of these creatures returning fo r
such distances when l iberated in regions remote from thei r
,
T he M a s o n
B ees
55
1914
2 78

I
habitat H e cannot attribute thei r return to m em o ry for
he s ays : I t was certainly not m e m ory but some spe cial

faculty wh ich w e must content ourselve s w ith recogniz ing
by i ts astoni shing e ffects w ithout p retending to explain it
s o greatly does i t transcend our own p sychology
B ethe thinks that bees returning to the nest

fo llow a
fo rce which i s entirely unknown to us and which causes
them to return to the place i n s pace fr om which they ew

B uttel Reepe n tri es to show that B ethe s exper iments fur
n ish such excellent proo f o f the existence o f memory for
locati o n that w e can hardly w i sh fo r anything bette r and

i n a footnote to hi s paper he shows that Fabre s unknown
sense theory w a s abol i shed a s early as 1 8 9 5 by some ex p eri
ments by We is smann who nds that the o nly co rrect solu

tion o f the enigma o f path n din g by Chali eo d om a i s that
the insects nd t hei r way back w ith the ir eye s

S o per
sistent i s Fabre i n cl i nging to hi s unknown sense theory that
again i n hi s chapter on red ants he asks why i f they i ssue

from the sam e mold has one a sense wh ich the othe r has
not an additional s ense constituting a much more over
powe ring factor than structural details I w ill wait until
evolutionists give me a val i d reason

Thus one can see how ever ready some natural ists are to
attribute any unusual behav ior t o an additional sense o r
some unknown powe r
So fear ful are they o f being
branded as i ndividuals ready to attribute som e aspe ct o f
behavior to an anth rop o morphi c conception that they p re fe r
to call i n some unknown inuence to account fo r conditions
I n the chapter on H oming i n M ind i n An imal s M r
E M Sm ith says : At one tim e the return o f bees to the

hive or pigeons to the cote a fte r foraging or explorative
exh ibitions was attributed to insti nct and th is implied
that the act i s l ittl e short o f m i raculous and absolutely i n
1
B uttel R eep en T he N a tura l H is to y o f the H o n ey B ee p 26
'
1 0
'
28 0
the home
There i s as the experiments seem to show
a l imit to the distance from which they have the powe r to
return I f a wasp i s taken far enough away from home
it can be lost even though it be w ithin the range o f its
phys ical endurance Now i f they were endowed w ith some
myste rious power o r sixth sense or magic o r unknown
fo rce or what not they should return to the nest regardless
o f the di stance they h ave to traverse s o long as it i s eas ily
w ithi n thei r phys ical power Furthermo re s ince P olis tes
do not return at night as Sm i th says then the suggested
unknown powe r i s p roved o f known impotence Again i f
the mysterious s ixth sense we re adequate w e should see
all o f the wasps d rawn as i f by mag ic at the sam e tim e to
the nest and w e would not nd such v ast variations in the
time for thei r return as f rom twenty two minutes to two
and one
fourth days Thi s unknown force o f B ethe and
othe rs shoul d transcend time and space but our ex p eri
ments th roughout leave every i nd ication that age experience
memory and perseve rance are the factors that bring P o lis tes
p a llip es back to thei r homes and when they are removed
beyond the di stance w ithi n which these faculties are ade
quate they are lo st Thei r case i s i n f act quite anal ogous
to that o f human be ings who when lacking these factors
o f age and experience p e rseverance and memory are easily
lost
.
'
P o lis tes
rub i g i n os us
L ept
We have seen these wasp s out on various occas i ons

from April 2 to October 1 3 and yet we have only a few
desultory stories out o f thei r whole l i fe h istory
One Ap ri l morn i ng whil e looking down from a clay bank
into a pond ten fee t below w e saw a P rub igi n os us a t
tempting to alight on the sur fac e o f the stale water We
,
28 1
have o ften seen them stand in the midst o f a body o f wat e r

without any disaster to drink While intently watching
he r alight w e were startled to hear a shri ll shriek as a
frog f rom under the bank pounced upon her and d i s ap
r
e
a
d
e
under the water The frog had been at rest on the
p
shore twelve inche s f rom the s p o t where the was p sought
anchorage and he j um p ed captured her shrieked and
dived all in les s time than it takes to tell It all went s o
quickly that one could not tell whether the frog shrieked
be fore o r a fter it captured the wasp but anyone w ho might
have heard i t would agre e w ith us that there w a s agony
almost human in that scream and we d o not think w e are
asking too much license in letting our fancy explain the
case f o r us
At Meramec H ighlands on August 1 7 1 9 1 5 we saw a
P
r ub ig i n os us
behaving queerly on the ground
We
watched i t fo r a few s econds when suddenly she pounced
upon a green caterp illar on the unde r s ide o f a lea f ab o ve
he r head i nvi s i b le to us b ut clearly w ithin her vi ew from
her pos ition on the ground S he carried it to the upper
side o f the lea f and began to malaxate i t all along its body
for several seconds and then continued to bite chew and
eat its anterior portion A fter she had eaten about one
fth o f the caterpillar s he again practiced malaxation on
the remainder ; then a fte r resting quietly on the lea f for
some m inutes s he ew away A fter s he had eaten her ll
and had taken the pains to knead the remainder we were
surpri sed that s he did not carry i t to the nest
On a sunny a fternoon at the very end o f the wasp season
October I 3 a numbe r o f P r ub i g i n os us and P a n n ul ar i s
were observed on the south si de o f the ol d rock ice house
at M eramec H ighlands I t seemed as though they were at
play ying to and fro against the build ing resting a whi le
and occasionally chasing one anothe r
Most o f them
,
WA S P S TUDI E S AF I EL D
2 82
seemed larger than usual What surp rised ! is very much

was to s ee s o many at leisure for one never sees a P o li s tes
but i t i s either on the nest or nervously itting from bush
to bush i n quest o f p rey and at th is date they ought to be
in h ibernation
We watched these P oli s tes perhap s twenty ve o f them
for about three hours and deci ded f rom thei r manne r that
the warm sunshine had lured them from thei r hibernating
places and that they were en j oy i ng some activity be fore
agai n entering thei r cold gloomy holes
The di lapidated stone building had numerous crevice s b e
tween the stones but most o f them were small ; a ve ry
few were large enough to pe rm it the wasps to enter and di s
appear f rom v ie w
The wasps were ying against the
wall in various place s exam ining a number o f crevices in
rap id succession all much too small
O ften they would
y away for some distance wheel around and return and
eventual ly ente r one that w a s large enough and remain
therein
P resently w e s a w that soon al l o f them w ould be gone
so
one by one w e took e ighteen o f them for study I t i s
a strange fact that o f these eve ry one o f the P a n n ula ri s
were males and all o f the P r ub ig i n os us were queens We
have o ften wondered i f thi s condition w a s the same
throughout the ock
They continued to play sometimes only one on the
wal l and sometimes as many as s ix until all had found
sheltering crevices o r had been captured All seemed to
have the same end i n view to nd a suitable lodging place
The enti re per fo rmance w a s con ned to the sunny s ide o f
the building although the o the r wall s o ffered eve ry induce
ment excepting the sunshine
O f the e ighteen which were taken home and kept in the
labo ratory only ten w e re l iv i ng a week late r They all
.
'
2 84
wa sp s T hey were resting or ying idly a round among

the rocks and a lo n g the sunny railroad track below ; by far
the greatest numbers were to be seen in the sheltering
crev ices o f rocks i n the cl i ff Care ful examination o f the
swarms p roved that the wasps were all o f the one species
al l queens and most interesting o f all po ints tha t they

were al l gregarious W e d i d not on that day nd a s ingle
sol itary individual M any o f the group s were small f rom
three to ten queens and others were larger ; w e estimated
three colonies to contain from twenty ve to fty wasps
each and one about a hundred resting quietly together
wh ile a fou rth colony was enormous w e do not think an

estimate o f a thousand would be excessive Even the mem
be rs o f thi s large group were s o docile that we coul d eas ily
approach near enough to them to ascertai n the sex At the
s ide o f the stream bel o w many were drinking water I t
was o f c o urse too early in the season for nest building
but we w ere elated w ith the thought that a l ittle later i n the
season w e shoul d nd a wonde r ful supply o f material for
observing how they begin to build thei r nests
A few dead wasps we re found scattered about the ledges
and frequently when w e chipp ed o ff a piece o f loose rock
a dead queen was to be found in the cranny behind it
I n two spots there wa s evi dence that this el iminati on
had been particularly heavy O n th e ledge behind two
colon ies o f mason bees were a great many w ings o f these
insects but no bodies These two sites we re p robably good
places fo r a n n ularis to spend the w inte r and als o good
places fo r a l izard a mol e o r a bat to come and feed It
was evident that the wasp s had not died a natural d eath
s ince only the w ings remained Whatever may have been
thei r f ate i t seem s that the i r severe stin g has avai led n o th
i ng in giving them protection
Later in the season June 2 5 we returned to thi s place
.
'
285
wi th v isions o f wondrous wasp activities

Imagine our
surp ri se when not one P a n n ular is was to be seen and no
nests on the rocks as we had expected Only tw o P oli s tes
were seen that d ay and they were P p a llip es We c anno t
account for this strange disappearance unless they had
dispersed or migrated
S everal li zards were seen along
the railroad track at the foot o f the blu ff but w e cannot
believe that they could ann ihilate so large a population o f
insects wh ich are so well equipped for ight and de fense
Yet again on this date i n one crevice i n the blu ff w hich
seemed an ideal h ibernating noo k we found upwards o f
fty queen s o f P a n n ula ris dead Thi s shows that we
must count upon heavy el imination
The interest o f thi s April congregation o f P a n n ularis
was only increased when on Octobe r 2 5 1 9 1 6 we found
an assemblage o f one hundred or more queens behind a
i
cl o sed shutte r o f an old abandoned building at Cl i fton Ter
race I llinoi s Thi s particular window wa s the most shel
t ere d nook on the building sunny and well protected from
the raw winds We searched at all the other shutters
but n o wasp s were behind them Whethe r the love o f com
p a n i o n s hip or the mere attraction o f phys ical c o m fort had
drawn them thus t o gether cann o t be declared ; we only
kn o w that they all had the abil i ty to c o ngregate in the
warmest spo t and n o t one was to be found else w here
in that vicinity The gr o up compri sed three species o f
P o li s tes ; perhaps seventy v e per cent o f them were p al
lip es twenty per cent a n n ula ri s and ve per cent b elli c os us
It appeared that the enti re populati o n o f the neighbo rhood
was here assembled yet there was not a male among them
At Raleigh North Carolina B rimley has taken males
all w inte r f rom the end o f N o vember to the end o f Mar c h ;
but while we have lo o ked for males duri ng the winter we
.
1 2
1 2
Ent N e
.
: 1 07
1 908
2 86
WA S P S TU DI E S AFI ELD
have neve r succeeded i n nding them I t seems as though

i t i s normal f or the m ales to di e o f exhausti on after f ertil iz
ing the queens but i t m ight be possible fo r s o m e un f o r
tun at e ones w ith no opportunity t o function s exually to
retai n su f ci ent vital ity to prolong thei r longevity
Whil e al l species o f P olis tes customarily place thei r nests
in buildings w e found s everal very large last year s nests
o n shrubs in a small
uncultivated patch o f ground about
fty by o n e hundred feet i n t he Ill inoi s valley ( S ee g
Wheat eld s surrounded it on all si des The nests
were attached to the bushe s about fou r o r ve feet from
the ground The striking feature was that several o f these
old nests had a number o f queens o f P an n ularis clustering
upon them I t w a s Apri l 2 4 1 9 1 4 j ust the time i n the
spring when the wasp queens would be l ikely to go house
hunting They may have been only resting on these old
nests but w e have long had a notion and so have other
w rite rs t hat they sometimes use o ld nests and that mo re
than one queen sometimes nds a home on such a nest ;
hence w e accepted this evidence i n partial conrmation o f
our imp ress ions The following table show s the number o f
queens that were at rest on each nest :
.
No queens on each
N o nests
o
I
5
6
1
th is w e found several new nests i n the making

o f three o r four cell s each some containing eggs and p re
s ided over by queens a s follows :
B esi des
2 88

Nests with
Nests w ith
Nes ts with
queen
queens
queens
nest
nest
nests
The re we re also some queens in c lusters w ithout any evi

dence whateve r o f nests being built
That ol d nests are som etimes used but usually not the
ol d cell s i s evi denced by the fact that ourishing P o lis tes
nests are o ften found w i th bright clean cells along the
periphery and a center o f soiled partly disintegrated cell s
H owever one should not attach to this fact too much
assumption o f intell igent choice fo r w e o ften n d nests in
strange s ituation s which are p robably accidental as for
instance the mud nests o f Cha l y b i on and P el op o eus o ften
found plaste red upon P olis tes nests or l ike one curiosity
which we s a w a p ipe o rgan nest and two mud nests all
subj o in ing a large paper nest Whether the queen fall s
upon old nests as accidentally as s he does upon the ra fters
o f a barn or ot her s ite i s a p roblem worth study ; but
w heth er through luck or foresight we must admit that the
ueen
enlarging
an
ol
d
nest
w
ith
the
stem
and
foundation
q
al ready built and the material at hand which may be again
used ha s m uc h o f her work al ready done at a time o f year
when s he i s w ithout as si stants
We usual ly think o f P o lis tes as i nhabitants o f sheds and
i n such places they are most abundant but thei r nests a re
built i n trees as well where the re i s no shelter whatever
f rom w ind and rain Judging by the large size o f these
nests which o f course denotes strong colonies we s ee that
they are little a ffected by the elements
They seem moreover to be able to adapt the i r nesting
habits to unusual conditions We once found a nest in a
rusty tom at o c a n i n a city rubbish hea p P o lis tes v ar i a nts

,
'
1 3
H un
g f
er o r
an d
W i ll ia m s
Ent
N ew s
231
25 5
91 2
2 89
in Kansas W hen without trees and buildings suspend the ir

nests i n the tunnel s o f rodents
The nests o f P an n ula ris sometimes attain great s ize
The foll o w ing data recorded by P ierce a re o f interest :

O f two nests taken i n Texas one had 1 5 7 5 cells and the
,
FI G
52
n es t o f
P o li s tes
a n n u l ari s a m o n g
t h e b ra n c h e s
other had 1 2 1 2 cells These 2 78 7 cells gave forth 1 5 5 3

wasps o f which 1 3 1 1 were males and 2 4 2 we re females
Two hundred sixty six wasps we re stylopized by Acros c his
m us p alli dus B rues
H e says further that stylopized P
a n n ula ri s a re not s o active a s normal i ndividual s
but are
o ften s een feeding on owers and some l ived f rom 1 3 to
.
'
14
B ull
N a t M us
.
No
66
: 17
1 909
2 90
days a fte r the exit of the male p a ras ites f rom the ir
5
bodies S ome were even found hibernating thus Fox
found at Denis o n Texas a nest o f P a n n ularis wh ich
measured eleven by sixteen in c hes
The large nest o f P an n ul aris illustrated i n gure 5 2
was found overhanging a dry creek bed near S t Louis It
was almost on edge among the b ranches eight feet f rom
t he ground and bore about twenty queens
They were all
f rightened away by the removal o f s o me o f the leaves in
o rder that the nest mi ght be photographed The nest had
p robably been built elsewhere and e ithe r the w ind or h igh
water had lodged i t h ere i n an unusual pos ition app a rently
w ithout having caused any change in the l i fe o f the progeny
I n one c ase w e n o ticed a stran g e phenomenon : one cell
i n a P a n n ula ris nest contained two eggs The second egg
was additional not merely a m isplaced one for the 2 8

cell s o f the n est contained 2 9 eggs Ten days late r w e
noticed however that only one egg in this cell was develop
ing ; the other seemed not to have hatched
One P a nn ulari s returned seve ral times to our out door
dinner table i n the c o unt ry to eat the j uic e f rom a bowl o f
appl e s auce
I t drank long and steadily stand i ng knee
deep in the j uice
M i ss M urt f elt r ecords the fact that these wasp s removed
strips o f paper from th e bags that were used to protect
clusters o f grapes A fter chewing the material into pulp
they carried i t away for the build i ng o f the nest P i e rce
2
loc
cit
p
2
nds
that
thi
s
w
asp
i
s
the
host
o
f
th
ree
)
(
species o f stylop s Rob ertson mentions it a s feed ing u p on
var i ous o wer heads and we have o bserved it f eed i n g u p on
owe rs o f the madeira v i ne and goldenro d
1
1 5
En t
N ew s
57
1 896
2 92
th is stone to observe the beh avior o f t he queen taking

p recauti o ns however to lay anothe r b road stone o f the
sam e color ab ove it i n orde r to give he r the roo f a n d the
shelter from the l ight an d weather to which s he w a s accus
to m ed
S he remained on the upturned nest until 3 p m
about ve hours and then deserted it The next day the
l i ttle black ants were steal ing the goodies from the exposed
nest Wat ch was kept at fr equent intervals for the next
two days but the queen neve r returned
Anothe r small nest o f P pa l lip es bui lt on the sheltered
s ide o f a loose doo r wh ich leaned again st the s ide o f a barn
was exposed by turn ing the door to the di rect rays o f the
August sun The nest w a s so o n dese rted by the owners
S hortly a fterward we found t hat the larvae too had been
to rn out p robably by bi rds
Al l wasps are usually s o di ligent about thei r domestic
duties that we seldom catch them do i ng s o commonplace a
thing as eating i n fact ; we are almost surpr i sed when w e

nd t hem pausing to take nouri shment fo r themselves
Early on e August morning w e found a P p a llip es vi s i t
ing a small wound i n a w i llow tree I t remained for per
ha p s a hal f hour w ith i ts head bur i ed i n th e so ft m o ist
ti ssues but wh en i t le ft w e could s ee that i t carr i ed away
n o w oo d p ul p Henc e w e concluded it was only break fast
ing o ff the j u ice and p ulp
A cottonwood tree bo re a decayed spot on the top o f a
p roj ecting root T w o P p allip es were busying themselves
about i t when w e di scovered i t at
a m Th i nking it
woul d be a good chance to see them gathe r pul p for n id i
cati on we watched the i r every movem ent A f ew m i n
ut es later a black and y ellow Eumen id came t o the sam e
place as i f returning to a fam i liar spo t Dur i ng the next
hou r whil e w e remained the y came and went at i nterval s
o ften paus i ng on the rotted wood f or ten m i nut es or mo re
,
. .
293
When they departed w e could see clearly that they never

carried away any pulp They spent the i r time w ith their
heads buried in t he rotted wood w ith thei r abdomens pul
sating in and out w hich movement in the wasp generally
denote s contentment We thought that they might be
me rely seeking moistu re s o w e placed near them a p iece
o f j uicy peach f rom our lunch This they stol idly ignored
how ever although they s eemed to be awar e o f i t an d even
cl imbed ove r i t on thei r w ay to the much p re ferred rotten
pulp H ence w e again concluded that i nstead o f gathering
the wood pulp f or the con s truction o f their n ests they were
merely enj oying thei r own repast
They are not always vegetable feede rs however On a
nest hung above the laboratory d oo r o n e worker brought in
a caterpi llar A second immediately cooperated w ith the
rst and together they s oon div ided it int o tw o equal parts ;
then each t o ok hi s portion and complacently chewed it
to pulp
1
We have prev iously recorded how when one comes
upon a P olis tes nest at ni ght one nds the i nhabitants
quietly at rest the i r b od ies an d legs sp read at against the
un de r sur face o f the nest They seem to be fully asleep
fo r one may hold a strong l ight nea r them fo r several min
utes be fore they S how the slightest response Early last
year when the queens were nesting al one w e wondered
w h a t b ecam e o f them at n ight
It se em ed that t hey went
el sewhere to sleep until a care ful ex a mination o f ftee n
ne sts o f P p al lip es revealed them in hi ding during the
night on top o f t he nest and completely invisible from b e
lo w
There they clun g to the stem o f the nest between
thei r roo f and the ceil ing o f the barn Thi s condition w a s
constant fo r all the queens at that stage o f n idi c a tio n and
,
16
An n E n t S o c Am er 9
.
: 24 1
916
2 94
brings to l ight the interesting vari at i on o f pos i tion in sleep

when the queen i s alone and when the numbers are great
D r C H Turner ha s published a very interesting and
detai led account o f the f eeding homing and hunting b e
18
h a v i o r o f a n or p han c olony o f P p allip es T o wer has
done some work on the ontogeny o f the colo r patte rn i n
thi s wasp P ierce nds P p allip es stylopi zed by the tw i st
Ac ros c hzs m us b ozvdi c hi and says that a
ed w ing parasite
femal e w a s taken by M r Dury so laden w ith paras ites
that it could not mo ve

I t contained nine mal e pupa e
Robe rtson nds them feeding on various ow e r heads
P ackard says says that a mal e with the abdomen removed
l ived fo r ve and one hal f hour s whil e a femal e w ith the
0
head removed l ived actively fo r fo rty one hou rs S nyde r
says that i n the B e rmuda I slands P p a llip es i s o f a l ighte r
S hade than in the Uni ted S tate s B ermuda has coral roads
and white c a ls o m in ed roo f s and he suggests that s elective
el imination has been a facto r in causing the surv ival o f the
l ight colored i ndividual
.
1 7
'
19
P oli s tes b elli c os us
Cres s [ S A Rohwe r ]
.
Tw o specimens o f P oli s tes b ellic os ns a yellow striped

wasp and als o anothe r Hymenopteron o ften cam e down to
the wate r to d rink Occasionally they stopped at the water s
edge but mo re o ft en they would y to the m i ddl e o f the
puddle o f still water and there al ight upon the sur face
spread thei r legs in the fashion o f the water stride r and
drink long and deep apparently in full enj o yment T hev
-
1 7
1 8
P s y c he
D e c em
P ub l
P sy c h e
2 : 1
1 9
20
E nt
News
: 1
84 - 1 90
U n iv C hi c a g o
1912
I
: 1
87 7
47
1 9 08
1 0 : 21
Fig
2
7 74
1 9 03
2 96
had lost hi s h ead at the choppin g b lock the h ead w a s

fou nd teem ing w ith thes e greedy sc avengers They j o s tled
and swarmed over the tongue eyes and a ll exposed mo ist
sur faces ( g
In early spring the queens always keep close to the
ground in ight and o ften ente r open ings like m ole holes
o r crevices i n the rocks We could easily understand thei r
l o w ight and hunting behavi or since it i s well known that
th i s s pecies bu ilds pa p e r nests underground ; the y were prob
ably not fo raging but nest seeking o r s ite hunting
On
May I 1 1 9 1 5 w e saw many in the elds at M eramec H igh
lands conducting themselves in thi s manner
-
FI G
m o v ed
53
Ves p u la
r o o s t er s
g er m a n i ca
hea d
ere
as
thi c
as
ie s
on
f res hly
re
Whi le they usually build thei r nests underground we

once f ound one in a hollow log lying on the ground It w a s
,
297
discovered early in j une and then comprised three empty

cells s ix covered ones and about a dozen w it h larvae F ro m
the covered cells four adults emerged on June 2 8 and 2 9
and July 3 The nest w a s small and had four laye rs o f
covering over the top and partly overlapp ing on the sides
,
Ves p a
ma c ul a ta
A pai r o f i nsects tumbl ing v i olently about o n the door

step attra c ted o ur attention Closer observation revealed
Ves p a m am la ta i n deadly combat with a large gray y S o on
one w ing o f the y ew o ff ; then the head popped o ff
Even then more tumbling ensued until presently the wasp
triumphantly adj usted the y under he r bod y and ew to
the grape a rbor nearby H ere s he paused to com plete the
preparation o f he r p rey wh ich consi sted o f much malaxa
tion o f the y and b iting o ff i ts other w i ng ; then S he ew
away w i th i t
For days t herea fte r S he chased ies at the kitchen door
and at ou r out doo r dinner table Other wasps were o ften
d rinking our fruit j uice ste a l
there too sharing our food
ing our cake even pillaging ou r sausa g e but Ves p a; ma m

la ta neve r bothere d anything but the ies which con gre
gated there W ith a swi ft ness exceeding that o f a chicken
hawk i n its onset upon a barn yard s he would appear above
the table pounce upon an un suspecting y e ither on the
w i ng or resting on the tabl e and carry it up to the grape
arbor overhead There S he w ould eithe r snip o ff its wings
and head p reparatory to carrying it away o r as is her
custom hang hersel f up by one hind leg and swing to and
fro up s ide d own while devouring the prey
They are large boi sterous and decidedly te rr i fying in
thei r manner
Howeve r they themselve s do not always
,
'
2 98
escape for w e once found one dead ensnared in a spi der s

web
Wasp s o f all kinds are alm o st always a hot weather
population ; hence w e were much surp r i sed on November
1 6 t o nd a V mac ul a ta
very active upon the window
pane I t w as l e ft there that its voluntary course o f action
m ight be observed That day was unusually warm ; on the
cooler days the wasp was only S l ightly active On Novem
be r 2 8 i t was placed in a j elly glass containing co tton but
e ither the c onnement d i d not suit i t o r its days were near
an end f or i t surv i ved less than two days m
ore
In January 1 9 1 0 a large n est w a s t aken at Kimm sw i ck
M issour i hav i ng several dead was p s lying between the tiers
o f the paper
9
9
3
FI G
54
n a tura l s i
T h e turret
o ver
the b u rr o
Ody n erus g em i n us
of
T wic e
ab o ut o n e fourth i nch i n diamete r and two inches in t o tal

length
It goes straight downward for hal f its length
then turns abruptly usually form i ng an angle o f ap p ro x i
mately 60 w ith the perp endi cular although occas ionally
th i s angle appro aches a ri ght angl e The end o f thi s c hannel
i s very S l ightly w ide r than the upper po rt i on and f orms one
o f the pockets wh ich make up t he nest ( s ee g

Other channel s w ith a S im ilar pocket may branch o ff f rom
the mai n burrow at the p o int o f the S harp bend
We
have found from one to four pockets to a S ingle nest but
w e do not know whethe r th e wasp s vary so i n thei r h a bits
o f nest building or whethe r the smal ler nests were only in
compl ete
We suspect that the constructi o n o f a s ingle
cluster o f cell s extends ove r a consi derabl e length o f t ime
because i n o n e nest o f four pockets wh ich we excavated
one cell conta i ned a larva o f conside rabl e s iz e whil e th e
others contained smaller larvae o r egg and caterpillars
1
H unger fo rd and Will iams nd no mud tub es over the tun
nel s o f O g emi n us and a part i f n ot all o f the pellets o f
earth are depos ited w i th in tw o or three i nches o f the en
trance to t he hol e We nd that O g emi n us drops he r
-
'
En t
N ew s
23 : 25 3
91 2
T HE
EU M EN I DAE
1
9
3
p ellets neare r to the en tra n c e tha n does 0 d ors alis the

latte r scattering them over a much larger area We too
have seen an occasional nest o f g em i n us W ithout the turret
but we attributed these cas es to som e accident the in tru
sion o f man o r beast We w atched three such nests whose
turrets had been destroyed to see whet her they would be re
constructed but no attempt w a s ever made by the wasps
to repai r them
FI G
m k
ar
55
s on
P h o to g ra p h
t he
a ll o f
the
O g e m i n us s ho i n g the ma n d ib l e
N a tura l s i e
v e rti c a l c ha n n e l
f the
n est o f
O f course such a pi ece o f w o rk as this pe r fect little

chimney would be impossible w ithout mud whereas the y
are usually built in dry barren places H ence this wasp
l ike her s i ster 0 d ors al is carrie s water and di sgorges it
a l ittl e at a time to wet the s o il making j ust enough mud
at a time fo r one pellet
We have never been s o fo rtunate as to w itness the begin
ning o f thi s pretty to wer but we have seen the builder
carryi ng out the pellets o f mud throu gh the chimney and
,
2
0
3
dro p p ing them a few inche s f rom the burrow : The aecom
p anying gu re ( g 5 6 ) illustrates a numbe r o f these
pellets w hich n icely S how the marks le ft by the manipula
tion o f the mandibles
We suspected th at the di rt taken out o f the hole w a s
appl i ed to the chimney but we were perplexe d that the
wasp S houl d apply part o f her excavated clay to the tur
ret and w ith the same care ful p reci sion carry out and
d iscard other pell ets
Finally near the end o f the season
on S eptember 1 6 w e got a cl ue to the answer to the ques
tion We found a turret which se emed incom p lete ; i t went
straight up f o r one th ird inch and had n o curve or ho ri
z o n t a l portion
The wasp was carrying out mo i st pellets
but i nstead o f applying them to the turret s he w a s taking
them out a few inches from the hole and d i scard ing them
When w e returned the next a fternoon the turret Stoo d at
p recisely the height at wh ich we had le ft it The wasp
w a s no longe r carrying out pellets but she emerged eve ry
f ew m i nutes and played about the mouth o f the ch i mney
Imag i ne our surp ri se when cl o ser scrutiny revealed to us
that she was at these times b iting o ff a mouth ful o f c l a v
f rom the m argin o f the c himney a fter hav i n g moi sten ed
the spot w ith water j ust a s she does i n digging her hole
and carry i ng i t d o w n i nto the burrow ! H er work was
me thodical and accurate really cha rm i ng to see
S he
woul d come up to the top o f the turret spread a drop o f
water b ite o ff and knead together a large mouth ful and
carry i t down hea d rst into the hole and come up in a
moment to repeat the p er formance p recisely The fact that
S he mo i stened only enough clay at each trip to make the
desi red chunk indicate s aga i n that s he must carry the sup p ly
o f water i n some other w a y than i n the mout h ; perhaps
deep down i n her throat When he r su p ply o f water w a s
R e turn i ng a fte r a
exhausted S he ew away for more
.
0
3 4
down below S u re enough when all the turret w a s gone it

w a s proved that the material i n the chimney w a s exactly the
amount requ i red to ll and seal up the burrow ush w ith
the s ur face o f the ground H ere then w a s our explanation :
the wasp digs he r burrow and carr i es out the ea rt h in moi st
pellets ; all the d i rt fr o m the gallery she care fully saves and
construct s i nto a chimney whil e that f rom the chamber o r
pocket s he carrie s out and discards When the nest i s pro
vi si oned s he plugs up the mout h o f the c h ambe r and pack s
back i nt o the gallery the ve ry same clay which S he took
out and has at hand exactly the requi red amount o f l le r
and al so mat erial o f the r i ght color to rende r the plug in
her hole i ndi stingui shable from the surrounding earth B y
this method the securi ty o f the young i s well sa feguarded
al so for the ent i re channel i s s o rmly packed that i t i s
impos sible to probe i t or even to trace it when digging as
one can o ften easily do w ith th e nests o f the wasp s that
kick t he dry di rt loosely into thei r burrows
Thi s wasp w o rked arduously in packing down the last di rt
on the top o f the hole moi sten ing it w ith the remainder o f
he r load o f wate r A fter s he had ni shed applying m o re
earth s he fetched one more load o f water and appl ied it
to the plug p resumably to make it more rm and compact
S he stood pounding and smoo thing the 3seal down w ith
her head her body curl ed almost in a ci rcl e so her abdomen
nearly touc hed her head and he r folded w i ngs st i cking
up vertically
Thus i t seems wasps vi e w i th one anothe r in wonde r ful
way s o f e ff ecting th e secur i ty o f the i r young and each n ew
method seems more marve llous than the last
An othe r turret building Eumen i d and i ts l ittle chimney
came unde r our notice at about 2 p m The ne st under
o
n
round
could
not
have
b
e
en
an
extens
ive
e
as
could
be
g
seen by the p au c ity o f the pelle ts strewn nearby so when
.
'
'
THE
EU M E NI DAE
9
3
we s a w the wasp carrying i n cate rp illars w e suspected

that i t w as her rst cell that S he w a s pr o vis ioning
The top s oil in that place w as black loam to a depth o f
one fourth inch while the subsoil w a s o f red clay T he
basal hal f o f th e turret w a s made o f black earth and the
uppe r hal f o f red clay and all o f the pellets strewn about
were o f red clay
This color arrangement p roves that
the wasp begi ns immediately upon commencing her bur
row to construct he r turret out o f the rst soi l excavated
and adds the de ep e r mouth ful s o f c lay in thei r turn hi gher
on the turret until the required size i s atta i ned ( or the re
quired amount o f material reserved ) and throws away the
remain ing pellets taken out o f the chamber or such po r
tions o f the nest as w ill not need to be re lled w i th earth
Thi s chimney w a s o f the standard f o rm and d imensions
The prop r i etress w a s car rying in caterpillars The rst
one w a s pr o cured in ve minutes and the second one in
twenty v e minutes
Wh il e she wa s gone w e made a scratch
on the su r face o f the ground to help us to locate the nest
Thi s con fused or alarmed he r so that S he ew about for
several m inutes be fo re entering Finally s he ventured into
the nest but ew out again and away car rying he r cater
pillar w ith her Not until a hal f hour had elapsed d i d s he
return and then S he c a m e empty handed Once more she
entered her hole and ew out again uneasily and began
to dismember the turret pellet by pellet and carry the clay
i nt o the hole We have n o t ed previously how they p lug up
thei r holes by removing all o f the turret and ca rrying it
i n for lle r so w e thought that S he w a s n ow do i ng the
same although we we re surpr i sed that she should n allv
seal up the t unnel when apparently only one cell w a s n
When about one fourth o f her chimney had
i s hed below
been taken down w e caught her to make sure o f her iden
tity and p roceeded t o o p en the burrow G reat w a s o ur s ur
,
9
3
pr i se upon nding that s he had placed a plug o f mud i n the

mouth o f t he cell that she had lled w ith caterp illars and
another thin layer ove r the top o f the hole ush w i th the
sur f ace o f th e g round while t he tunnel which fo r p erma
nent seal ing i s always completely lled w a s empty H e re
w e s ee that when danger threatened and insuf cient t i me
was at hand to seal the bu rrow normally S he d id th e best
th i ng p ossible t o s afegua rd th e youn g i n the shorte s t p o s
s ible t i me
S ince we o ften nd from two to four cell s
e manat i ng i n all di rect ions f rom the central tunnel i t might
be that i n th i s case since thi s nest had only one cel l her
intentions were to return and construct ot her branch gal
l eri es and cells from the bottom o f th is em p t y main channel
when S he felt that an i ntruder w a s not lu rking near S uch
behav i or ought to make man feel l i ke a vai n p retender
wh en h e presumes t o lay an exclus i ve cla i m to intelli gence
The burrow w as the regula r ve rt i cal tunnel w i th the
chambe r at the bottom turn ing to one s i de as w e have a l
ready phot ographed ( s ee g
There we re seven cater
p illars in the chamber o f the spec i es o f Lox os teg e d i s
cussed ful ly elsewhere They were all act i vely mov i ng and
coul d even walk about
The egg w a s hang i ng by a t i ny thread f r o m the wall I t
had been depo s i ted at 3 p m on S eptember 1 0 and i t
hatched at
a m on S eptember 1 3 thus hav i ng a per i od
o f i ncubation o f about two and one hal f days The m i n i a
ture la rva w r i ggled out thr o ugh o n e end o f i ts shell and fell
right down among the squ i rm i ng caterp i llars Thei r a c tiv
i ty seemed to do i t no harm ho w ever for it thr i ved for ten
days and became large and husky until one morn i ng i t
was found dead Fabre th i nks that the egg woul d be i n
j ured by th e wr i ggl i ng mass o f cate rp i llars i f it we re not
hung up out o f the i r reach but here w e th i nk as w i th O
dors alis the egg mi ght be placed among the cat erp i llars
,
0
8
3
In the vicinity o f the tu rret j ust described w e found in

the bare ground some plugs o f mud sealing former holes
These were no longer a mystery to us a fter we had seen 0
h
e r c himney
break
down
m
a
k
e
m
ud
o
f
i
t
an d
i
u
e
m
n
s
g
,
FI G
57
s t ruc ti o n
B urro w s
N atura l
O dy n erus g em i n us in
of
si e
v a ri o u s
s tag e s
of
c on
clo se th e hole ush with the sur face N ear each o f these
were strewn twelve o r fteen ol d pellets o f mud o f the type
wh ich g em i n us always throw s a w ay when excavat i ng her
cham be r The rst nes t w hich w e dug out seem ed a p e r
f ec t p iece o f work Clustered around the base o f the vert i cal
.
T HE
EU M EN I DAE
9
3 9
channel were four pockets al l al ike and all co ntaining cater

pi ll a rs o f the s pecies used exclusively by O g emi n us ( See
gure in lower right hand corner o f g

There were
f ty tw o c a t erp i a rs in all or an averag e o f thi rt e en to a
cell They were writhing bri skly an d se em ed ve ry much
alive b ut t hey w ere unable to w alk The larvae we re smal l
but appar ently healthy An o ther o f the sealed b urrow s near
b y had pr eci sely the sam e external ap p e arance b ut had
only one cell below ground ( uppe r right hand gure ) This
contained eleven cate rpillars o f the same spe cie s o f L ox o
s teg e as the others and the egg was hanging from the wall
by a short but strong s ilken thread
O ne certain incomplete turret surmounting an o p en hole
was w atched for th ree days b ut it n either i ncreased nor de
creased in height and S inc e we saw no S i gns o f activ i ty
a b o ut it w e d ug i t up The m ain c hannel went d own as
usual fo r an inch where it branched and t ermi n a ted in
two p ockets ( s ee g 5 7 lower le ft gure ) One o f these
c o n tained a large 0 g em i n us larva and was well sealed
with a plug o f mud in the mo uth o f th e cham b er whi le the
second w a s open and contained only four caterpillars and
apparently was unnished P robably the mother was p had
m et w ith an acc ident which had prevented he r f rom n i sh
ing the nest but the incomplete nest gives u s the evidence
that the turret i s l e ft standing unt i l the p ocket s be low are
a l l nished
H unge r ford and Wi lliams have some evidence th at the
cells are o ften us ed more than once as brood cham b ers
They also illustrate man y galleri es and cells to one nes t
apparently m ade by one mother The most com p lex nes t
i n o ur experience i s that shown in gure 5 7 i n which the
cells diverge f r o m the central gal lery in a sh o we r
When we o p ened one new n est We were s ur p r i sed to nd
i n its single c ell one lone cate rpillar and the mother w a sp
,
WA S P S TUDI E S
1
0
3
A FI ELD
Th is burrow w a s dug out at ab out 5 p

the
little own e r h ad m erely gone into her burrow for a n ight s
l o dg i ng o r i t m ay be th a t w e had only caught her there
when busy w ith n i d i c at ion
While w e sus p ect that O
i
e
m
n us sleeps i n he r hol e the sam e a s do es 0 d o rs alis w e
g
fou nd o n e ins ect aslee p on M eli lo tus cl ing i ng to the stalk
H oweve r the s ex o f thi s i ndividual w a s not noted
The v e ry rst 0 g em i n us w o rker o f the y ear was seen
i n the eld on Jun e 5 but they were no t at al l common
unti l Augus t and mo st o f the ab ove notes w er e mad e i n
the rst hal f o f S eptember
Som e wh i ch w e reared at
home eme rg ed a s adults in June O n September 2 4 w e
not ed that du r i ng that w eek the tu rrets o f g emi n us ha d
been scarce Th i s d isap p ea ran ce o f w asps w a s proba b ly
due to the i ncreasing coolness and the fact that thei r no r
mal s eason w as ove r ; but i t may als o h av e b een due to the
fact that al l the puddle s o f w ate r i n the vici nity h ad dri ed
up i n the long continued drought
O f the hundreds o f c a te rpi llars exhumed f ro m th e ne sts
o f O g em i n us every one w a s o f the sam e s p ecie s They
w ere i denti ed by D r S B P rack er as belong i n g to the
genus L ox os teg e o f the fami ly P y ram tt da e Dr Fracker
writ es fu rther that i t was di fcult to be sure o f the s p e cies
since the alc o hol had t aken o ut all the color but i n al l
probabi lity th ey w ere L s i m i l arlis Thus the two s p eci es o f
O dy n em s OI g em i n us and O d o rs a lis dwelt togethe r i n
th i s eld ( g 2 ) and found plenti ful food for the ir young
yet each adhe red strictly to her o w n ch o ice o f pr ey the
on
e using only L ox os teg e and the othe r taki ng exclus i vely
P holi s ora c a tullus
S i nce the tw o s p ec i es are s o S imi lar i t
seem s to us su rpr i s i n g th at they should b e so strict in thei r
cho ice o f prey as n ev er to acc ept what i s enti rely pleasin g
to the other H un g e r ford a n d Williams n o te a simila r
.
'
'
'
En t
N ew s
23
25 4
W A S P STUDI E S AFI ELD
2
1
3
sta n d a t once w hy i t m i ght be sa fer fo r the delicate egg

to be fastened up to the wall by a strong thread than to be
dropped r uthlessly i nto a w r i thing m a ss like thi s although
we sus p ect th a t i n m o st c ase s it would surviv e
A few o f the cater p illars showed excep tion al longev i ty
Tw o remained active for f ty four d ays and one aston
i s hed us by continuing to l ive f rom S eptem b e r to the fol
low ing April or s even and a hal f months After the rst
thre e mo nths i t ceased spontaneous w rithing and moved
only upon stimulati on but remained plum p and healthy
looking unti l the last Doe s the p o i son o f the st i ng act as
?
a pre servative
I t must have som e such po ten c y f o r w ith
out i t und er natu ral conditions these ca te rpi llars i m p ri s
o n ed W i thout food
woul d have died and decayed i n a day
or two o r pupated b ut i t seems that a fte r the wasps have
ni shed w i th th em they can do n e i the r
Ody ne rus d ors alis
Fab [ S A Rohwer ]
.
Each S p ec i es o f wa sp has its ow n highly s p ecial ized

method o f building its ne st Some bite out the earth i n
chunks some di g i t to pieces an d carry i t out in arm fuls
others scratch and k ick it out accord i ng to various xed
h a bits S o 0 d ors um to o h as her own p eculiar method
which di ffers f rom the others She c arries water mouth ful
by mouth ful and mo i stens the spot o f hard dry earth so
converting i t i nto mud which S he carries out i n p ell ets We
have never chanced to see he r working except when the
earth wa s d ry and hard ; w e S hould like m uch to k now
whethe r she continues to carry water when she di gs he r
nest afte r a ra i n P erhaps th ei r method o f wo rk may well
be gle a ned f rom the d etail ed description o f the behavior o f
a S ingle i ndiv i dual wh ich seemed to u s typical in p ra c ti
cal ly all points
.
1
3
When w e d i scovered t hi s wasp at wor k a t 8 :5 0 on the

mo rn ing o f July 3 1 her h o le was al ready begun and a bout
one hal f the l ength o f her body in depth She would stand
on he r head in the hol e and bite out a l ittl e clod at the
same time turning around screw l ike a b out one qu a rter
or one hal f a revolution in order to work f ro m al l d i rec
t i on s al ike Then she would ba ck out o f the h o l e with her
round well formed pellet o f mud i n her mandibles alw ays
y f rom two to fteen inches w i th it an d dr o p it These
mud balls were fai rly uni f o rm in si ze and round not mere
clods o f di rt bitten out ( Se e the p ellets strewn about i n
'
g 58 )
.
A fter working thus for a b out ten minutes S he went away

p robably fo r a f r esh supply o f w ater We waited so long
for h er return that w e began to think that s he had ab a n
d o ned the enterpri s e ; but a fte r fte en m inutes she re a p
a
e
S he worked almo s t
p red and began her wo rk a f resh
inc ess antly and too k o ut 1 5 pel lets i n 8 m inu tes Wh en
s he em e rg ed fro m the hole w ith th e la s t pellet and ew
a way a f ew i nches to drop i t she did not retur n i m m ed i a t elv
to continue he r burro w ing but cont i nued her i ght at a
hi g h speed and i n a d i rect line toward the northwest She
wa s gone f rom ou r si ght j ust one and a hal f m inutes when
w e s aw he r returning straight from the same d i r e ction
3
I sel ey s a ys that w hen they ret u rn from thei r waterin g
places he can s ee the i r mouth
parts gli stenin g ; we were un
able to s ee s o much but s he resumed her wo rk as be fore so
we suppose s he carried the water in he r gullet Thi s time s he
w orked 8 m i nutes t he same a s be fo re and i n that p eri od
took o ut 1 2 pellets ; upo n em erging to cast aw ay the last
j ust as
S he ew strai ght and sw i ftly o ff to th e no rthwest
Thus S he seem ed
be fo re and returned i n TM m inutes
to wo rk w ith pretty p recisi on and ac curacy yet w e doubt
1913
Ka n U n iv S c i B ull 8 : 2 87
,
1
3
FI G
a te r
59
Th e
c o u rs e o f
lg h t
of
O dy n er us d o rs a li s t o
th e p ud d l e o f
pel lets when S he soa red o ff to the northeast W e waited fo r

her at he r o ld pon d and although S h e had own i n th e op p o
s ite di rection s he eventually ap p eared there p a us ed a m o
m ent for a l o ad o f w ater an d hurried on back to he r hole
follow ing her habitual route preci sely and ar riving a fter
an absence o f j ust 6 % m inutes Why these round ab out
co urses ? We d o not know but thi s show s that even though
these wasps are fam iliar w ith the di rect route they do not
always adhere to it The hole w a s now about two inches
deep ; the pellets were larger and fewe r to each mouth ful o f
wate r Eithe r the wo rk w a s becoming more a rduous or it
was tel ling on her strength fo r she m oved more heavily
and occasionally stagge red when s he d ropped a ball
The
ne x
t three w orking periods were 8 6 and 5 minutes and in
them she took out only 8 9 and 9 pell ets Our St Loui s
wasps generally wo rked at a d iff erent pace f rom I s el ey s
K ansas 0 d ors a li s ; the latte r made the trip for wate r in less
than a mi nute but w ith each load o f water they removed
only 5 or 6 pellets o f earth wh ile ours always took out f rom
to
I
pellets
7
5
I n making he r tri p tow ard the n o rthwest she now simpl i
e d her cours e s omewhat to a more n ea rly d irect route
simi lar to the ri g ht hand diagram in gure 5 9 Once more
.
TH E EUM E NI DAE
1
3
S he w ent o ff on her northeasterly t o ur and w a s gone 1 5

minutesin fact so long th a t s he had trouble in l o cating
,
her burrow P revi ously s he h ad com e to i t d i rectly ev ery

time witho ut a m
oment s con fusi on but a s unhesitatingly as
a man returns to hi s own hom e
S he ew ab out as i f
con fused w ith i n t w elve o r ei ghteen inches o f the bur row
for over a minute then S he gave up the search w en t b ack
to the weeds about twenty v e feet a way I n the d irecti on
from which S he usual ly cam e and from there c ame back to
i t di rectly show ing again t he necessity o f g oin g bac k to her
starting point for he r bearings I t w a s then necessary for
us to go When we le ft her she was c o nt i nuin g her north
west trips by the o val route last described I n desertin g thi s
wasp we probably mi ssed a most excellent opportunity o f
for thi s nest was p robably
see i n g the com p lete n i d i c a ti on
nearing com pleti o n
.
FI G 60
.
T he b u rro
of
O dy n er us d ors a li s
N a tura l
si
The nests are always ab o ut two

two and one hal f
inches in depth and a quart er to th ree ei ghths inch in di am
or
-
1
3
r one qua rter
ter at the t op The hole goes strai ght down f o
to one hal f an i nch and then w i den s to f o rm a s p her i cal

chamber a hal f inch i n diamete r ( s ee gs 60 6 1
F rom the
b ottom o f th i s the gallery continu es the s a m e d iameter as
be fore for a sh o rt di stance and expands into a s econd ch am
b er S im i lar t o the one above
The l ower c ell i s lle d w ith
cater p i llars the egg i s le ft sw inging by a del ic ate thread
f rom the wall o f the cell and the mouth o f the com p artment
i s seale d up w ith a plug o f mud ; then the upp e r cell i s s im i
l a rly lled and seal ed w i th a mud plug i n the neck o f the cell
Finally a closure o f mud i s i nse rt ed i n the to p o f the h o le
ush w i th the s ur face sometim es art fully construct ed s o
but
a s to be i nd istin gu ishable from the surround i n g earth
more o ften th ere i s the sauce r like de p ress i on caused by bit
i ng out ea rt h to ll the hol e w hi ch I s el ey h as s o w el l de
scri bed B ut ri g ht here i s a very i ntere st i ng and cur i ous
point : the entire gallery from the top o f the u p p e r cell to
the sur fac e plug i s n ot lled i n sol i d fo r i ts ent i re length as
a re the holes o f othe r bur row i ng wasps although o ften i t
i s a sho rt c hannel ( g 6 1 ) but i t has only the rm plug
at the top and bottom and an a i r chamber between The
purpose o f th i s ar rangement leads one i nto pretty spec ul a
t i on Does th e a i r chambe r i n the channel help to ma i nta i n
?
T he
b
n
c
a
a more un i fo rm t em p e ratu re i n the ells e e th
depth i s somet i m e s so sl i ght that the d i ff eren ce could not
amount to much at any event
I seley too nds that the n est s are vert i cal w i th one o r
H e desc ri bes one exc ept i on howe ver wh i ch i s
tw o cell s
v ery s u rpr i s i n g and i nterest i ng : a colony o f e i g ht nests
bu i lt i n the fac e o f a ve rt i cal c l ay bank w i th from t hree
to seven cells to e ach bu rrow We have n eve r yet fo und
m o re than two cell sexcept i n on e nest which w as appar
ently abnormal which had an extra ( small ) cell b etween the
two normal ones ( gu re 62 ) ne i ther have w e found these
e
2
0
3
I n all o f the channels and nests may b e seen on the si de

walls the m a ndi ble m ark s i n late ral strokes o r hal f rings
where bites were taken out ho rizontal ly not up and d own
The gure als o betrays the m ethod o f pro
( See g
c ed ure in nest diggin g : as the len g th o f the burrow i n
c reases the w idth also increases so that a t the b ottom there
i s a s ta rt on the rst gl obular cel l
When thi s has bee n
c o mpleted and provisi oned and the egg d ep os ited undoubt
ed ly the w idening o f the sec o nd cell occurs
and perh aps
the soi l remov ed in thi s wi d en i n g g o es to fo rm the plug
for the lo wer cell
When th e home i s at last i n ord er and M adam Ody n erus
goes a mark eting she does pr ec i sely what m o st othe r wasps
sc ru p ulously avo i d : S he l eaves her nest w id e o p en T o b e
sure the re i s yet n othing i n i t to attract o r reward marau
de rs b ut ne i ther I S there i n the newly n i s hed bur row s o f
other s p ec i es so one i s again l e ft wondering w hat m ay have
been the o r i g i n o f such an i nstinct what condit i on can

have brou g ht such pressure upon th e s p ec i es as to estab
l ish i n t hem so xed a habi t
I seley ( l oc c i t p 2 8 9 ) w as ve ry fo rtun a te to observe
her hunt i ng in a patch o f mallow
W hen 0 d ors alis
come s upon a crumpled l ea f c onta i n i n g the larva o f the
s p o tted S k i p p e r she commences tearing energeti cally at the
si lken nest rst at one end and then at the other Some
times mo re than ve m inute s i s r equired to d i sl od g e a
cate rp i ll a r but more o ften the v i ctim i s j erk ed f rom i ts
cove r i n less tha n a m inute s ei zed by the n eck and stung
tw o o r th ree tim es under the tho rax V i g orou s mal axat i on
follows the stinging a fter which the caterp i llar is carr i ed
to the nest w i thout del ay
These w asps when return i ng f ro m the eld se em to
experience no d i f culty whatever i n l oc a t i n g t h e i r bur
rows They car ry the i r p rey o n the w i ng u sual ly w i th com
,
THE
FI G
62
An
a
p ra ti v e
un u s ua l
b u rro
EUM E NI DAE
of
O dy n e rus d ors a li s
2
1
3
N a tura l
Si
ease hu gging it ti ghtly all the while alight at the

brink o f the hole and push i t in ahead o f them holding to
i t as they lower it ( see g
then they f o llow it int o
the hole remai n inside for a few seconds perhaps a hal f
m inute and then come backing out S o m etimes the wasp

soars away d i rectly i n quest o f other game ; at othe r times
she S its down a n d washes her fac e fo r a moment then rises
and poises on vibratin g w ings an inch or tw o abov e the
hole turns around on the wing as i f ins p ecting the s it e
then ci rc les abo ut and ie s away She i s calm gentle and
com p o sed in all the maneuvers b etrays no n ervou snes s and
wastes no time i n blustering
We had lon g suspected that thi s w asp builds not o n e
burrow but a group of burro w s and w ere glad to read
I seley s statem ent o f hav i ng seen one begin a second hole
,
2
2
3
a fter the rst Was ni shed and close beside i t

On several occas i ons when w e b egan to dig up a nest and
scooped o ff the sur face soil with the trowel the removal
i m m ed a tely
FI G
r
T he m a n di b l e m a
63
N a tura l S i e
.
on
the tun n el
of
Ody n er us d o rs a l is
the thi n laye r revealed three or four nests near togethe r

We hav e only occasionally noticed in ou r St Louis wasp
eld the pretty sauce r l ike depressions which betray the
location o f the cl osed ne s ts
One such group o f nests particularly i m pressed us Fo r
a week at lea st we had been watchin g a certain O d o rs a li s
as we crossed the eld each morning On every occasion
o
2
3 4
not o vi p osit at the conclusion o f the provisioning as do

othe r species but whenever i t suits her fancy o r phys i
olo g y We sometime s nd the little egg s w in g in g from t he
w al l on its thread when the provi sioning i s o nly b egun or
hal f done
Among the St Loui s po pulation o f d ors alis we have
found only the one specie s o f caterpi llar used a s prey vi a
the H esperid la rva P ho lis om c a tullus [ Drs B arne s and
M c D un n o ug h ] while I seley has found tw o s p ecies o f L epi
d o p tero us larvae used by t he Kansas 0 d ors al is the s p ecies
menti oned a b ove and P j rg us tes relma
We a re not p ositive that O d ors a lis always sleeps i n her
bur rows but we have found her i n the unni shed holes at
twi light and on dark cloudy days a su fc ien t num b e r o f
t imes to m ak e us cert ai n that at least th e females d o so
frequently whi l e the di g g i ng i s i n progress
O f course
the problem s remain o f whe re the males sleep and where
the female s nd S helter when they have not open holes at
thei r di spo sal
I t has always seem ed to us rem a rkable t h a t the nests o f
O d o rs a li s are not more o ften ravaged b y parasites o r en e
m i es since the owne r always leave s the burrows wide o pe n
durin g her long absences Yet w e believe S he su ffe rs le ss
at the hands o f such impostors ( at least in the nests which
have c ome und er ou r notice ) than do so me o f her s iste r
speci es wh ich always close thei r burrows behi nd them w ith
such exact i ng care
F rom on e open hole w e s a w an O d ors a lis emerging
and watched her closely to as certain he r method o f egress
Im a gine our surprise when we s aw he r ca rry out a P c o tul
lus caterp illar in her f ront and m i ddle pai rs o f legs pause
at the b rink o f the hol e for s everal s econds to adj ust the
We
n ev er
ha v e
y et
c a tullus.
o b s e rv e d
ha v e w e
f o un d
d ors a li s in th ree
h er
us i n
c a terp
w id
el y
il la rs
s ep a a t e d
o th e r
tha n
a r ea s
b ut
P ho l is o ra
TH E EU M ENI DAE
2
3 5
caterpi llar pro p erly between forelegs and mandibles and

y a w a y wi th it Was s he a deliberate robbe r or was she
only a home seeker w ho had by erro r entered the wrong
nest and w a s removing only that which S he had brou g ht
?
h
These wasps are
w i t her and which w a s right fully hers
usually very care ful i n nding thei r nests and while they
sometime s pe ep into two or three others b e fore a rr1 v1 n g at
thei r own w e have seen only thi s one go so far as to ente r
the hole o f anothe r
One day we sa w a black w as p running and hopp ing about
i n the r egi on o f the holes o f O d ors al is fo r abo ut ve m in
ute s ; nally it enter ed one o f the holes and we c aptured it
as it em erged to ascertai n its identity I t w a s a N o tog on i
d ea a rg en tat a E ve [ S A Rohwe r ]
In th is case too w e
w ere uncertain wh ether N o tog on i d ea was invading the nest
o f Ody n erus through a blunder or through m a l intent Thi s
w asp hunts crickets and s he probably entered thi s hol e i n
search o f them
Certai n Diptera sometime s pester Ody n erus One two
celled ne st which we found appa r ently securely sealed
contained ei ght D i pte ra pupae
On the sam e day on whi ch we made the ab o ve discovery
S eptembe r 1 8 w e w atched a small gray Dipteron H i lar ella
n
[ C H T Townsend ] following an O d o rs al is w hich
wa s homeward b ound with her green cat erpi llar The l itt le
y tagged behind her m ost p e rsi stently keeping j ust at a
sa fe distance a few inches to the rear o f the was p I t fol
lowed and p oised in the ai r w ith all t he ski ll which insects
o f thei r pro fession usually di splay B oth ew near to the
ground a nd i f 0 d o rs al is rest ed for an in stant the y
would hove r or poi se on v ibrating wings a f ew inches away ;
i f the rest was p rolonged the y wo uld rest on the ground
or a grass bl ade nearby I t w a s pretty to see the shadow
on the white n ed earth o f the two insects in the bright sun
,
2
6
3
moving i n uni son as i f they w ere i nf i s ibly united

The w a sp ew h ithe r and thither i n an evas iv e w ay in the
regi on o f her nest fo r fully tw enty minute s be fore nally
ente ring he r hole We have never Se en another o f these
w a s p s e x p e ri en c e such di ffi culty i n loc a ting h er nest ; can i t
b e that thi s on e s p ent thi s time and effort yin g to and fro
?
i n an e ffort to evade her pursuer
We have neve r w i tnessed t he attack o f the was p upo n the
caterpi llar b ut as I seley says they are stung and paralyzed
T hey k eep fresh plump and g re en and usually respo nd to
stimulus w ri the and voi d excrement fo r from thre e days
~
to a w eek a ft er thei r entombment ; and w e hav e had s o me
Whieh conti nued t o l ive and rem ai n plump for two o r three
weeks o r even more F ollowing are som e typical r eco rds
w hi ch show the longevi ty o f the prey ( all cate rpi llars mov
ing actively about when taken f rom the nest )
5light ,
'
'
'
Nest
Nes t B
N est
died a fte r 8 days

1
died
a
fter
days
3
5
1 died a fter 2 5 days
1 died a fter 1 2 days
1 d i ed a fter 1 4 days
I d ied a fte r 1 9 days
I died a fte r 2 0 days
days
1 di ed a fter
3
2 d i ed a ft er
6 days
2 died a fte r
days
9
2 died a fte r 1 0 d a ys
I d i ed a fter 1 4 days
I d ied a fte r 2 2 days
1 died after 2 6 days
I died a fter 2 9 days
I
N est
I d i ed
N est E
I
1
1
N est P
a f t er 2
day
s
5
die d a fte r 2 7 days
di ed a fte r 6 d ays
died a fte r 8 day s
died a fter 1 4 days
d i ed a f ter 3 days
died a fter 5 days
di ed a fte r 6 d ays
d ied a fte r 9 days
di ed a fte r 1 2 days
died a fte r 1 5 days
d ied a ft er 1 8 days
'
I
I
1
1
I
T he
n um
n um b e r o
b er
f day s
of
d ay s
s in c e
f ter
th e i r
th e
c ap
w
x hum d
m wh t x
d
ne s t s
tur e
so
e re
c ee
Of
th i s
c o u rs e
t he
2
8
3
another not only sucking out the j uices but c o n suming the
enti re carcass excepting the dry shell o f the l ittle black
heads ( s ee g 6 4 the remains o f a f east ) The supply o f
food provided by the mother fo r the gro wing in fant varies
greatly but we have neve r asce rtai n ed whether that su p ply
i s sometimes actually inadequate o r excessive o r wheth er
the young me rely accepts whateve r i s g iven i t and get s
through som ehow I n our e a gerness t o kee p al ive all tha t
w e had at hom e for observati o n we gave all o f them addi
t i o n a l fo o d and it was almost always accepted greedily
Once we w ere s o daring a s to o ffe r a fat p rospe rous larva
a small brown caterpilla r instead o f the customa ry green
ones T he larva tu rned at onc e to thi s new fresh food
but mumbled about i t m incingly for only a m inute and then
went back to resum e chewin g at i ts old h a l f eate n gr een car
cass We suspect that s o m e o f t hem are i n real need o f
more food b e fo re they pupate especi ally sinc e s ome h av e as
We
f ew as three little caterpil lars upon which to subsist
chanced to nd on e goo d
si zed larva wh ich must have been
eating di rt fo r the a b domen w a s full o f earth Yet over
feeding probably tends to mak e them o ve rf at and delicate
We had one whi ch w as a thriving i n fant and prom ised to
be a ne adult A fte r it had suc ked dry seven caterpillars
i t was enormously large and f at AS w e trans ferred i t to
a cle a n b o ttle in the hope o f see in g it spi n its cocoo n we
let the vial fal l to the table and th e j ar caused the bo dy
wall s o f the fatling to burst O f course i n nature the larv a
runs no risk o f such a catastrophe and yet the ci rcumstance
sug g ests that over feeding may make them a s well as othe r
organi sms so ft
There i s an astoni shing d egree o f variation in the dura
tion o f the di fferent p eri od s o f d evelopment in thi s s p ecies
There seems to be no xed time for hatchin g spinning o r
any o f the functions M oreover w e have not yet a s c er
,
T HE
ta i n e d
EUM E NI DAE
2
3 9
any causes climatic envi ronmental or he ditary to

account for such variations Thi s variabil ity w i ll be easi ly
noted from the tab ulato n o f a few typical cases as foll o ws :
,
D at e
o zri
'
F i n i s hed
wn
H a tc hed
os i t
9
9
10
h
t
5
1 1
2d
h
t
5
9
8 31
d
3
d
3
1 1
6 th
6 th
10
h
t
5
day
day
day
day
day
eed i n g
S p un
th
day
4
1 5 th day
8 th
day
inj
ured
(
1 0 th day
1
day
day
day
day
1 5 th day
1 8 th day
1 5 th day
1 5 th day
1
th
day
4
day
1 4 th day
1
4 th
4 th
On August
w e exhumed a three fourths grown larva
with its last caterpillar We gave i t three more which i t
e a gerly devoured A fte r ve days more i t b egan to spin
I t was unable to construct i ts c o co o n normally in the tin
box ; s o w e made i n the box a nest o f tiny bit s o f so ft paper
After that i t succeeded be tte r i n covering itsel f but we
could still see the ins ect w ithin No t until August 1 9 or
twelve d ays a fter rst s p i n m n g did i t tran s fo rm into a
mummy l ike pupa with in the w eb O n September 1 2 a fte r
a pupal p eri o d o f thi rty six days an adult female emerged
There i s a p o ss ibi lity that the trans f ormation o f thi s i ndi
vidual may have been delayed by the impeded construc tion
o f the cocoon Thi s adult w a s kept in a cage and l ived on
sugar water fo r eight days T hi s t o o w o uld indicate that
at least two generation s emerge each year
One o f the perplexing p roblems is the question o f how
the wasp in the lower cell when adult can emer g e w ithout
To be sure
n jurin g its younger brothe r in the ce l l above
they probably l ie dormant all winter and are ready for
2
o
33
eme rgence at approximately the same tim e i n the following

sprin g ; and yet we can not s ee how even then they c ould
make thei r way out w ithout o ften coming into di sastrous
collision Thi s m ay account i n part for the large numbe r
o f dead adults o f thi s speci es more than o f B n u b ilip en n is
wh i ch we h ave dug out i n the spr i ng We have never as
c e rta i n ed whether these d ead wasps a re t he youn
adults
g
which neve r f o und thei r way out to the li ght o f day o r
w asps o f the parent generati on wh ich may have died while
sleeping in thei r bur row s o r m ay have been entombed by
the mud o f heavy rains
The vari ati o n in t he p e r i ods o f d evelopment menti oned
above may help t o clear up thi s con ict o f the emerging
wasps or again i t m ay only add to the c on fusion Thi s
point w a s suggested t o u s by a certai n burrow an ex
c ep ti o n a l nest in t hi s vi cinity i n that i t had thre e cell s
( g
We dug i t up j ust a fter the to p cell ha d been closed
One would logically expect t o nd the o ccupa nt o f the
lowermost cell o f the tie r the largest b ut we were surprised
to nd the larva in the m iddle c ell la rge the bottom one
small an d the uppe rm o st sti ll o f c ou rse in the eg g sta g e
H ow thi s state o f a ffai rs could have com e about remains a
7
mystery
We wan ted to follow up th i s d evelopment b ut
the lowest larva died H oweve r w hen they arrived at the
stage o f pupation the m iddle or la rg est one S pun i ts cocoon
onl y three days earli er than the smallest one ( which was still
in the egg stage when we found it )
H ence w e m ay b e
li eve that the di ff erence in the date o f maturation o f these
and the intermedi a te one would h ave been even less
Fab re nds that in certain bees one sex hatches earli er
than th e ot her and the mother c a n control the s ex o f the
T he
t o v a ry
s ex
of
f ac t that the fo reg o in g ta b l e S ho w s the i n c ub a tio n o f the

f ro m 2 t o 6 d ay s m ay m ea n a c o rela ti o o f the p eri o d to
r
t h e d e vel o p i n g
l a rv a
eg g
th e
2
33
o f the prey 1 8 not mentioned ; i t mi ght be that she uses sub

terran ean caterpi llars s ince our note s S how that S he enters
numerous holes i n a foraging m anner
Robertson nds the adults frequenting m a ny s p eci es o f
owers
,
O dy n erus d es i g n a tus
.
Two o f these Eumen id wasps we re found fast aslee p on a

i
B ot h were curled
p g wee d at 4 :30 p m on August 2 8
a round lea f petioles
At 4 a m the foll ow ing day i t
rained and rai n continued to fal l unti l 1 0 o clo ck
At
0
:
9 3 i n the morning they w ere i n the i dentical places and
positions and d ri pping w et
The rain had not a roused
them to activ i ty i n the least ; on the contrary they were in
a kin d o f stupo r which rendered them very easy to pick up
They w ere not alone i n thei r m ise ry N earby were found
a num b e r o f o the r i nsects P elop oeus ca em en tari um May
beetles and others behaving similarly
-
Ody n em s p ed es tris S a us s
A
S
[
.
Rohwer ]
O n a clay bank ( s ee g
r i s i ng about s ix feet a b ove
the leve l o f the ground w e s aw on October 3 thi s Eu
m en i d entering and leaving an open hol e at i nte rvals A fter
catching he r w e dug up th e nest ; th i s was o f the exact
si ze o f the draw i ng in gure 6 5
Thi s wasp make s a gallery and f ro m time to time con
as they a re needed Si m i lar to
s truc t s c ham be r s beneath it
the manner o f P hi la n thus
except that the latter usually
builds them f rom th e s ides o f the gallery whereas thi s one
builds the gallery dro ps a branch o f it to a cham b e r con
t i n ue s the galle ry and makes an o ther chambe r there from
and s o on
,
THE
EUM E NI DAE
333
The rst cell that thi s one had made the o n e nearest the
ori ce contained a hal f grown larva and four green ish blue
cat erpi llars all o f o n e specie s o f G elec hi idae [ S B
Fracker ] ( the larva was too im m ature to det e rm ine the
species ) The next cell contained s ix spe cimens o f the same
Lepidopterous larva and hanging by a tiny thread from
,
FI G
65
T he
n es t o f
O dy n e r us p ed es tri s i n
a c l ay
b an
N a tu ra l
si
the wall was the egg The small p etiole like gallery that
led from the main gallery t o thi s w a s ti ghtly packe d with
soi l and the continuation o f the main c hannel in a p a ra llel
d i rection led us to think that a thi rd cell was about to be
ho ll owed o ut o f the end o f thi s gall ery One has not far to
seek to suspe ct that the earth that i s rem oved i n the di gg i n g
o f the n ew galle ry and cell i s used for the ll in g o f the
sh o rt neck o f the cell j ust provi sioned
The fat larva which w e captured in the rst cell ate all it
coul d nd and al so tw o caterpi llars tha t we gave i t from
the oth er cell The next day when we introduced a large
-
334
one we found i n the eld i t ma de a bra ve e ff o rt to e at thi s ;

but its strength p roved unequal to cop ing w ith i ts tough
S kin aud its wriggl ing activity Two days later the larva
di ed
N o details o f b i ological sign i cance could be found i n
the l it erature o n thi s w asp
Ody n erus
S a us s
o ra m i n a tus
[ S A Rohwer ]
The rst Ody nerus f ordm ma tus w ith which we became

acqua inted we re nesting i n a pil e o f o ld weathered logs
They are sol itary dwellers and it i s highly d o ub t f ul i f they
live i n colon i es ; th i s instance w as probably a c a s e o f a num
be r o f them nest i ng near t ogethe r merely bec aus e the sit e
a ff o rded the prope r condition s for each one
O ne wasp i n particular attra cted our a ttention S he had
he r burrow i n a log twelve inches f rom the ground Tw ice
S he ente red ; t hen w e crept c lo s e to s ee what S he w a s s o
myster i ously doin g i n her l ittle cra nny She had about a
hal f dozen caterpi llars and wa s arrangin g and pac kin g
them c los e together unti l they were forced so deep into the
hole that they were out o f si ght
Then S he l e ft and s o on
returned w ith a m oi s t pellet o f gray re clay and use d i t
She brought a
insi de the hole probably fo r a part ition
second load ; then w e took her to ascertai n her identity
Nearb y w ere other nests o f thi s kind ( see g
w hic h were S imilarly situated As w e watched one the
We are
Was p s beh av i or took a m o st remarkable form
c er tain that o ur presen ce did not caus e i t for w e Were
care ful n ot to i ntrude Without apparent cause s he carri ed
out four cate rpi llars one at a time and droppe d them in the
We followed and found one o f them ;
ro a d S ix f eet away
i t w a s E n arm o n i a s p o f the family T ortricidae [ S B
,
9
33
b urrow
replaced it w ith the others and conti nued for some

tim e a rranging and pa cking them
One day one o f these wasps carry ing a green caterpillar
under her body w a s seen t o enter one o f the holes in a log
A fte r all e ffor ts to dig i t out had proved vain we re
m oved the log from the pi le and w ith g reat e ffort s hook
the caterpi llar f rom the hol e I t was not dead but re
sp o n d ed to stimul i
On ano ther occas ion an O f ora m i n a tus
was ca rrying hom e under h er body a G n ori m os c hem a g al
lam o li d ag i n is [ S B F racke r ] while j ust b eneath the mouth
o f he r burrow i n the lo g w a s anothe r a Lox os teg e s p
p robably s i m i la lis [ S B F ra c k er] caught in a sp ider s w eb
where she had evi dently d ropped it
This old log heap fo rm ed an excellent hab i tat for the
longi co rn beetles and many o f them had at some time o r
anot he r taken advantage o f the condition s and made thei r
burrows there Fo r a long time we took i t for granted
that these wasps used the deserted hole s m ad e by these
power ful be etles fo r i t was almost incredible that s o del i
cate a l ittle creature could cut deep smooth holes in oaken
logs which broke one blade a fter another o f a heavy p ocket
kni fe B ut one day w e fo und on e su rely ca rvin g o ut her
rs t and soon back o ut

She
would
ente
r
head
o w n ho le
and drop a little load o f f resh s a w dust on the g round She
worked fai th fully and steadi ly but her b urrow was already
o t s e e her method o f cutting the
s o deep that w e could n
wood She proceeded qui etly as i f modestly unaware that
she was doing something truly marvel lous Thi s must
have been a b out the i r hole c utting tim e fo r on that day

I n the l ogs
July
we
found
a
dozen
similar
holes
all
(
newly cut although we d id not intercept any other wasps
at work on them
We have commented elsewhere upon the remarkable lack
o f dissem ination a m ong many wasps
G ene ration a fter
,
'
'
'
T HE
EUM ENI DAE
337
generation they remai n pers istently in the place where they

we re born and frequently an entire populatio n i n a local ity
i s exterm inated j ust b ecause o f thei r i nabi lity or unwillin g
ness to m ove from a chosen spot even when it appears
to us that mi gration would be very easy And so it w a s
w ith this pa rt icular g ro up o f O f ora m ina t us
It seems
they we re introduced into thi s local ity about eight years
ago at a tim e when the brick manu fac turin g company piled
several carloads o f logs i n the eld These logs have been
hauled away from time to time b y cartload s and u sed for
starting the kiln res I t i s po ssible that they were intro
duc ed i n some stage o f the i r development w ith the logs
and nding everything to thei r liking have occupied the
site generation a fter generation neve r going even to the
fence p osts or other w ooden structures near by As a resul t
o f th i s i nability to go even a little w ay from th e place o f
thei r bi rth thei r numbe rs have grow n le ss and less as the
logs have been used ; and now when the last load will have
gone to feed the ame s there seem s no ho p e o f anything
but extermination fo r th i s lot We have made dil igent
search a b o ut the region to nd even a few o f them taking
up the i r abo de i n so me more permanent structure but w ith
out S uccess I f like ou r B em b ix n ub i lip en n is and S eo lia
dub ia they c annot d i sseminate ove r di fferent pa rts o f a
restricted area how i s i t possible for them to cover di stinct
?
o r remo te areas
Hunger ford and Williams dug them out o f a stump i n
Decatur County Kansas and found the cells se p arated by
partitions o f mud
A fte r so recently deploring the p robabl e extermin ati o n
o f many o f these wasps on acc ount of thei r lack o f ada p ta
tion to other place s o f abode than the logs w e we re grati
ed to have di scove red i n another area at least one c as e i n
whi ch th is i nsect had used a hollow twig Thi s elderbe rry
,
FI G
z
s i e.
67
T he
n est o f
O dy n erus
o ra m i n a tus
in
a n e l d er s t e m
N a tura l
o
4
3
completed o r we re complet i ng the i r develop m ent at about

th i s time and we re ready to emerge When taken f rom thei r
co coons i n the tw i g and placed on cotton saturated w i th
sweetened wate r they i mmediately pa rtook eage rly o f the
n o ur i shment
Anoth er 0 f oram i na tus w a s seen o n e June day carry i ng
a da rk gray c a terp i l lar i nto an old lo g smoke house She
h i d i t i n a burrow i n the lowermo st log and then sealed
the hol e with m ud
I t w a s i m p oss i ble to i nvest i gate the nests o f th i s s p e c i es
to s ee whethe r they too s n er f rom the ravages o f para
s ites O n m ore than one occas i on however w e s aw Chry
C hrys is ( H ex ac hrys is ) i n tri ca te B r [ S A Roh
s i s bees
wer ] ying about the pil e o f wo o d where i n the wasps
nested as i f look i ng for the bur rows o f some host ; hence
w e should not be at al l surp r i sed i f they sometimes meddled
i n the homes o f O f ora m i n a tus also P i erce nds the
f emales to be the host o f the tw i sted w i ng parasite S trep
.
si
t
e
r
a
p
Robertson found 0 f ora m i na tm females to go from

ow e r to ower o f P en ts tem on la ew g altus S olon d and
turn ing to the base o f the tube c ut a ho le i n one Si de

w i th her sha rp j aws and inse rt her tongue t hen S h e cut a
hole in the othe r s ide and agai n i nserted he r t o n gue The
necta r i s lodged on each si de o f the base o f the steri le la
ment and the wasp S howed remarkabl e i ntell i gence i n m ak
i n g a hole on each s i de
W e s aw them repeatedly feeding
at the owe rs o f M elilo tus al b a and Robertso n r eco rds
vari ous other owers u p on wh ich they feed
.
'
Am i s troc erus f ulw p es S a us s [ S A Rohwe r ]

'
Thi s w a sp which w e had previously observed enterin g

hole s I n logs was seen at the edge o f a corn eld near
,
T HE
EU M E NI DAE
1
34
Creve Coeu r Lake diggin g up mouth fuls o f mud Late r

i n the day anothe r indiv i dual w a s seen at the same place
doing pr eci sely the same thing We supp os e i t w a s carry
i ng mud to m ake partitions and plug s i n i ts burrow some
S everal others were entering
w here p e rhap s i n wood
holes in old log s piled a sh o rt d i stance away Some o f them
were digging in the woo d One i n particular entered a
shallow hole an d with the b od y pa rtly protruding turned
i tsel f a round so that i t w a s inverted and c ut away the wood
at the ceil ing o f the burrow Hence we s ee that they do not
use o ld tunn els o f o the r in sects but m ine thei r own or
enlarge thos e already at hand
We fo und that thi s s p ecies m akes good use o f the old
cell s o f m ud daubing wasps They use these old cell s w ith
o ut m od i cat i on
lling them with cate rpilla rs fo r thei r
yo ung and then s ealing the a p erture w ith mud Ashmead
cites Walsh who sa w one nest bui lding i n a s p o ol
Not alone doe s thi s was p us e the tunnels i n wood w ith or
without m od ica tion ; s he als o di gs bu rrows o f her own In
a small c lay b ank protect ed Ove rhead by a po rch three
s p e c i mens o f thi s wasp w ere se en at work as early a s June
28
As the seas on advanc ed others ap p e ared
The i ntere stin g feature o f thi s bank was t hat its face
was completely riddled w ith the ol d tunnel s o f a min ing
b e e E n tec fmi a tamrea Say ( see g
any one o f which
would have made an ample dom icile fo r th is wasp Instead
o f appropr i ating one o f thes e however S he always built
he r own tunnel s i n the clay bank although in s o doing s he
o ften broke i nto the barrel shaped cells le ft by this bee
One o f these wasps w a s seen i n the morning b ri nging in a
ca terpi ll a r and i n the a fte rnoon seal in g the cell w ith a plug
o f w et earth The burro w when o pened was f ound to have
two cells separated by a p art i tion o f mud The tunnel w a s
,
o urn
A n im a l
B ehavi o r
: 2 7 -63
191 6
2
4
3
abo ut one fourth inc h i n diamete r and each o f the two

cells was th ree fourths i n c h long The p lug at the ori ce
was thicker than the partition M o st interesting o f all w a s
the f act that the last cell ha d broken int o an old bee cell
and this too had not been used but o n the contrary ha d
-
FI G
n i ty
68
f
Ab a n do n ed ho l es
re a d y
m a de ho m e s ,
i n te r s t i c e s to d i g h e r
own
of
the m i n i n
An c is tr o c erus
b urro
I g n o ri n g
f ulv ip es f o un d
b ee
the
op
sp ac e
p o rtu
i n the
partitioned o ff w i th mud The cell n earest the sur face

o f the ground contai ned seven cate rpillars all o f the same
si ze and species and the egg a ttached to the wall by the
usual short th read I t hatched w ithin the succeed ing twenty
four hours The caterp illars too were all al ive at that time
and had been lett i ng out s i lk
b een
344
inch i n length and the o ther thre e cel ls w e re o n e and one

hal f inche s i n total length
The n ext nest o pened wa s unl ike any o f the othe rs ; i t
had but a s ingle cell and no ai r cham b e r We had watched
the mother was p close th e burrow w i th m ud and ex p ected
to nd the egg quite new
B ut the larva w a s alre ady
hatched an d w a s beginning to pa rtake of i ts supp ly o f pro
vi s i ons ( th i rteen caterpillars i dentied by S B F ra c k er
as E x er tem a
Thi s i s S i gn icant i n S how ing that the
nest is not fully provi s i oned b e fore t he egg i s laid but the
egg i s deposited be fore o r during prov i s ion ing
She does not make h er burrow by k i cking out the so il
but like certa i n species o f Ody n erus she carries wate r
so ftens the earth i nto mud which S h e gathers into a pellet
carries out and di scards She enters he r hole head rst
gets her load backs out and ies backward fo r about S ix
re
i nches then drop s the load and
enters We once counted
that w ith one mouth ful o f wate r s he brought out n i ne
pell ets o f mud ; th e succeed i ng mouth ful made exactly the
same number Another took out six loads o f mud i n three
minutes
The holes are l e ft open durin g the propr i etor s absence
A large blue cuckoo be e once entered a cell and remained
with i n for several m inutes
Thus we see he r a versatile creature who can adapt her
sel f to various conditi ons w ith equal success
,
An c i s tr o c erus
tig ri s S aus s
[ S A Ro hwe r ]
.
A mud nest was found o n the outside o f a schoolho use at

L ake V iew Kansas The nest s eemed newly made and
from its app earance we thought that i t belong ed to a
Cha ly b i on or a S c elip hron
When w e brok e i t o p en ho w
-
THE
EUM ENI DAE
34 5
eve r we found that eac h long cell had two partitio ns mak
ing o f i t three cells eac h o f which was lined w ith a pap ery
substance
Most o f the pupae were inj ured by the j ar
when the n est fell so that only two came to maturity and
eme rged as adult A tig ris on August 5 and 6 1 9 1 6 The
caterpillars remaining i n the cells were so dried up that
thei r spe cies w a s i ndeterm inable
At rst i t seem ed possible that thi s Eumenid had made
he r own nest i nstead o f being a renter b ut later w e f ound
evi dence o f the cells h a ving been resealed ; so i n all p rob
abil ity it had been prev i ously occupied by o n e o f the mud
daubers
M rs S los s o n nds thi s i nsect to i nha b it the Alpine re
gion o f Mt Washingt on B ank s nds that i t frequents
the owers o f C ean o thus and Robertson rec o rds i t as hav
ing been taken while feed ing on var i ous ower heads
T w o specimens o f thi s speci es were taken dead f rom a
spider s web They l inge r unti l late i n the fal l ; on October
2 7 several o f them were out feeding upon the few remain
ing owers o f the goldenrod at Cl i fton Terrace Illinois
,
10
An c is troc erus
c o ra
S a us s
[S
A Ro hwer ]
.
A woody elder twi g o f the prev i ous season w a s picked up

from near the ground at Eureka M i ssouri on July 1 2
1 91 6
I t was kept indoors all w inte r and on Apri l 2 3 2 7
1917
fou r wasp s o f the above speci es and one cuckoo bee
C hrys i s ( Tetrac hry s is )
[ S A Rohwer ] emerged
the re f rom
The rst cell the b ottom o f which Wa s on a level w i th
the ground was o n e and a quarter inches i n lengt h and
sealed at the top Wi th a mud part i tion The cocoo n therein
,
1 0
Ent
N ew s
2 3 : 1 07
912
9
34
'
was ve ry li g ht tan a lm os t w hit e thi n and p ape ry The
l
second cel was three fou rths inch i n length unlled and
l ikewi se cl osed at the top w ith a wall o f mud A b ove thi s
was another c el l only one fourth inch i n len g th occupi ed
b y A Cap ra
The next cell one hal f i nch long contained
a dead la rva The remaining four cells measured one hal f
i nch or less and g a ve forth adult wasp s o r paras ites
1
Ashm e a d says on the authority o f Rev T W Fyles
that thi s was p prov is i ons its cells wi th larvae o f the larch
sa w y
N em a tus eri c hs on i i and i s economically valuable
destroying tineina geomet rina tortric ina p ry alin a and noc
tuina larvae i n great num b e rs
,
M o n o bi a q ua d ri d em L i nn
[ S A Rohwe r ]
.
The carpente r m ud wasp

M on o b ia q ua d rid em ha s
been w i dely kn own for years but l ittle has been noted on
its li f e history That it i s a mud wasp i s true ; that it i s a
carpente r wasp i s doubt ful s ince the author s wh o attribute
thi s h abi t to th is wasp bring fo rward no pos itive evidence
except that the nests are found i n w oo d bo res This po int
w il l bea r furthe r i nve stigation and unti l it is determined
som e more app ropriate name should be appl i ed to the in
sect
The f ew know n f acts o f i ts l i fe hi story may b e
summed up as i n the follow ing paragra p hs
Ashmead says that M q uad ri d em p rey s upo n large
cut worms which sh e carries to her cells These c ells are
made i n the old bu rrows o f the c arpenter bee Xy l o c op a
m rg i ni c a, by rst reno v ating the s ides o f the old tunnel
w ith a thin veneering o f clay and then divi ding the s p ace
i nto cells w i t h clay partitions w orkin g from the bottom
,
'
12
'
'
1 1
12
P s y c h e 7 : 77
P sy c he 7 : 77
896
896
'
8
34
them carr i ed i t aw ay j ust as the mud dau b ers do ? She stood

on her head a n d struggled w i th the sti ff mud a s i f only w i th
great e ffo rt could she rem o v e th e clay wh i le mud daub i ng
P elop oeus at the edge o f the i r w a tering places care fully
select the i r mud o f j ust the ri ght consistency and then
gathe r i t up with c ompa rat i ve e ase an d grace
A year later August 7 1 9 1 7 w e s aw th ree i nstances o f
the m ud carry i ng propensiti e s o f thi s i ngen i ous was p One
came f rom a far to the d i rt bank unde r th e po rch whe re w e
were loi tering mo i stened a spo t o f the e arth unti l i t b e
came mud which s he gathered i n a b i g ball and car ri ed
away A second one wh i ch had her nest i n a boa rd i n the
po rch above the clay bank w ent out f or a load o f water
came back and al ighted on the turret made by a b urrow in g
bee and a fter soaking i t ap prop r i ated a huge mouth ful o f
her neighbor s p retty dwell ing i n a manne r most nonchalant
A third thanks to our vandal ism had not s o fa r to go fo r
her part i tion i ng mud F o r about tw o days w e had been
not i cing th i s o n e enter i ng an a b andoned car p enter b ee hole
i n the j o i st and wer e eage r to s ee the i nte rnal struc ture o f
the burrow
W i th a poc ket kn i fe w e attempted to re
mo ve enough o f the wood to ex p ose the cells but a fte r
expending m uch energ y i n wh i ttling w e decided i t could not
be done Suddenly it dawne d upo n us that to leav e thi s
f resh cut i n the weather worn p orch pro p would s p ell the
d o om o f the few pr i v i leg es wh i ch w e enj oyed o f p rowl i n g
about people s p rem i ses Si nce ou r errat i c behav i o r w as al
ready regarded w i th grave susp ici on by the p opulace o f
thi s rusti c community Then w e remem be red th e sav i ng
g race o f protect i ve colorat i on We h asti ly g ather ed a f ew
hand fuls o f mud and besmeared the br i ght cuts to rende r
all the woodwork a un i fo rm gray col or The wasp con
t i n ued he r work howev er and p roted at once by her new
convenience Although S he had been go i ng el sewhere for
-
TH E EU M E NI DAE
34 9
her mud f o r ten days o r more S he at once changed her

course o f action and gathered thi s mud which we had j ust
placed near to he r door We have neve r seen another wasp
s o quick to alte r he r activ ity i n order to avail hersel f o f a
new and strange conven ience or economy
The seve ral nests wh i ch have been ex am ined i nternally
S howed much var i at i on i n the n estin g hab i ts and provi sion
ing Wh ile Comstock i ndicates that these wasps us e vertical
hole s in trees w e have found them to oc cur i n various
other Si tuat i ons
The rs t one ope ned w as i n the under Si de o f a po rch
oor With m uch e ff ort su f c i ent wood was cut away to
ex p o se three cell s o f a large tunnel that went into the j o ist
hori zontally The s ides o f the tunnel bore marks o f pre
not a lone o f thi s wasp but also o f the
v i o us part i t i ons
carpenter b ee ; hence i t seemed evident that thi s hole had
b een used several times by both the bee and the wasp
There w a s no venee r i ng o f mud such as Ashmead describes
In one cel l w a s a fat gr o w i ng larva w i th S ixteen large cate r
~
p i llars all i dent i ed by D r F ra ck er as E p ip a s c hiin ae
p robably E p ip as c hi a 3 17 The othe r cell conta i ned eighteen
cate rpi llars o f the sam e spec i e s Th i s seems i ndeed an as
ton i s hin g numbe r c ons i der i ng the i r s i ze to be stored for
a s i ngle i n fant
Anothe r mother which was utili zing an abandoned b ur
row i ng b e e hole i n the clay bank fo r i ts dom i cile was
caught i n he r task o f carry i ng i n caterpi llars and sealing
the cell The rst one S he brought i n fteen m inutes the
second o n e i n twenty ve m i nute s These large c a terp il
lars were ca rr i ed very de ftly unde r the body ; the lon g trunk
was not all ow ed to d angle awkwardly but o n e end was
clutched i n the mand i bles wh i le the other end w as c las p ed
close to the abdomen wi th the legs When next obse rved
at 1 1 o clock S he w as carrying i n either m ud or water and
,
9
35
seal ing up the hole ; the aperture w a s alread y clo sed and
s he was o ccupied w ith biting o ut mud alongs i de the burrow
and adding i t to th e plug Suddenly S he ew away and
returned i n ve m inutes w ithout mud but w ith water and
moi sten e d th e e a rth beside th e plug We ea g erly pressed
too cl o se to study he r method and s he ed Three mi nutes
late r S he returned bringing a pe llet o f m ud from a far and
applied it w ent out and got anothe r bal l i n the same w ay
and added that ; then w ith the surplus w ater i n her m o n th
s he soaked up m ore earth b es ide the hol e
dug i t out an d
plastered i t also ove r the closure Here both methods were
used : b r i ng i ng l n water and makin g mud near at hand and
carrying mud from a far In applying thi s mud S he re
vo lved about the hole tak ing every po siti on describing a
ci rcle A fte r two m inutes s he brought i n still another b all
o f m ud Fo r a few moments our attention w a s d rawn to
another l ittle wasp nearby and when w e turned back the
rst had torn out all her ca re fully constructed masonry
opened her s eal an d w as busy inside the burrow What
S he w a s d o i ng there we could not ascertain
but when we
returned to the scene at 4 p m the bu rrow was once mo re
n icely sealed A large f reshly made depressi on bes ide the
h ole gave ev i dence that much o f the material fo r the plug
had been gathered ri g ht on the prem i se s Upo n o peni ng
thi s nest we found i t had rst an ai r s pa c e o f one hal f
i nch depth then a partition and a littl e way b ehind thi s
the remain s o f another partiti o n the one evidentl y b roken

int o w hen S he demoli shed the f ront door Through poor
mani p ul ation in opening the nest we got the contents o f
t w o cell s togeth er which comp rised thi rty six cater p illars
o f the k ind already menti oned and an eg g fo r each c o m
r
m
n
Almost
all
the
caterpil
lars
spontaneously
ex
a
t
e
t
o
f
p
hib ited si g n s o f li fe
The eggs were not carelessly tucked
i n among the writhing caterpilla rs b ut in true Eumen id
.
'
2
35
entered the hole w i thout turning around An old b ee c o

coon s he pulled out bodily and cast aside thus H ere w e
s ee an additional ac tivity o f th i s wasp ; enlarging holes t o
make them suita b le fo r her habitati on
Thus far i n the observati ons w e had never seen one o f
these was p s actually cutt i ng wood but i n all of the i n s tances
menti oned above and a numbe r o f others they had me rely
utili zed t he old ho les o f carpenter be es or other b o rers
H enc e we we re grow i ng suspicious that these s o cal led car
n
r
e
t
e
wasp
s
are
not
wood
cutte
rs
at
all
At
thi
s
j
uncture
p
our surm ise was strengthened b y nding a nest i n the hol
low roller holder o f an old i ron bed p ost on a j unk heap
O f course w e could not o pe n up the nest to exam ine its
structure but w ith the forceps w e rem o ved th e caterp illars
s ix i n number
Two w ere o f the s p ecies mentioned here
to fore a n d four were o f tw o othe r spe cies not noted b e fore ;
a m i shap prevent ed the i r i dentication but this record will
S how th a t M q ua dri dem uses more than one species o f
prey
O n July 2 5 one o f these wasps w a s n oticed i n the out
doo r l a b o ratory where she had a p propriated an old nest
o f the P el op o eus mud dauber Thi s nest was lyin g on the
shel f and had been used also by a Tryp oxy l on c lava tum for
nidus The cell i n the t o p row o f the nest was empty and
o p en and M on o b i a had eno rmo usly enlarged it s openin g
I t already contai ned two c aterpi llars and the eg g w a s hang
ing o n its thread Thi s n icely show s that M q zta dri dens
do es not rst prov i s i on the nest and then ovi pos it as do
som e H ymenopt era but that she lays the egg be fore or
during provi sioning T hat night s he slept in the cel l w ith
he r fac e tow ard the openin g and by July 2 9 when I re
turned to the farm she had com pleted an d sealed the cell
The
B et ween August 5 and 1 5 one male was p eme rged
cell when opened contai ned no evidence o f a cocoon but
re-
'
TH E E UM E N I D A E
35 3
two crumpled light brown shedding skins remained one

resembling the p upa and the other more larva like This
i s the second instance o f the absence o f a c ocoon ; pe rhap s
th ey do not s p in one
The b ehavio r o f insects i n the event o f rain is a po int sel
dom noted but we once had the oppo rtunity o f watching
these under such circumstan ces From our place o f s helter
under the p o rch and be side thei r ne sts w e could observe
them to advant age T he rst to come home in the rai n w a s
ve ry S hy i n attem pting to ente r her nest and when w e pe r
s isted i n lingering near to see i f S he carried anything she
ew angr i ly away in all the rain S oon ano ther came in all
wet and with slow ight but when we attemp ted to cap ture
it it too wh eeled a bout and went out into the stor m After
ten minutes it returned all drenched in the mos t terri c
down p o ur went di rect to the hole and crept in During the
h o ur three others were seen to come home in the heavy rain
in the same manne r The car p ente r bees nesting in the
same plac e did not c ome home but a fter the shower m any
came i n
Further observations mu st S how whether thi s wasp uses
sal iva o r wate r for m aking her mortar and i f she ever d oes
the w ork o f excavating her ow n tunnel in w o od I f s he
actually ca rri es wate r it i s remarkable that she does not
gather the mud itsel f di r ectly f rom the water s edge unless
i ndeed the streams thereabout are all so sandy or gravelly
that there i s no mud a b out them and S he must carry water
to an e a rthy area like the spo t on the sandbar in F ox Creek
to get her plaste r At least one can hardly imagine streams
mudless to such an extent that the development o f this habit
would be forc ed Thus the evolution o f such a habit as this
remains a wonder and mystery but since the wasp is so apt
in adapting ce rtain conveniences to her needs in all pr ob
abil ity some day she or her descendants will be found using
m ore ec o nom ic al building or renovating methods
-
W A S P S TUDI E S
35 4
A FI ELD
The habit o f this i nsect o f car rying wate r to m o i s ten the

ea rth is indeed a wonder ful in stinct I t i s even m ore sim p le
for Ody n erus d ors a le to carry w ater to moi sten the earth
where s he digs her bu rrow than it i s for M on o bi a to m ake
a burrow in a log or bank or even appropri ate an old hole
go elsewhere f o r water an d carry i t to a di stant d ry spot in
the earth make the m ud a mouth ful at a time and carry it
bac k to her nest for part i tion s and plugs
That w o uld
combine the methods o f a water ca rri er wasp a mud daub
ing wasp and a wood cutter plus a renovator or enlarger
o f old b ur rows T h i s would be i ndeed the most co m p lex
li st o f accomplishments and one o f the most c omplicated
methods o f ne st building found among wasps
9
35
P eckham
One may s afely s ay to o that the work o f the

Europe an naturali sts w ill not greatly exc eed thi s number
and that approx i mately l ess than 2 00 species o f was p s have
anything l ike a complete report to the ir credit When we
compare thi s w ith the number o f known species o f Vesp i dae
up to 1 8 9 4 and Sphegidae to 1 8 9 7 ( d e Dalla To rre s Cata
wh ich I S approx i mately 99 00 w e
lo gus H y m en o p
get an i dea o f the vast amount o f work yet to be done
be fore w e dare attempt to constru ct a synthetic genetic
psycholo g y o f wasps
The s o cial and sol i tary wasps a fford e s pe c i ally ne ma
teri a l wi th which to w ork out such pr o blems as the c o rrela
tion o f hab i t to structure th e o rigi n o f s oc i al ization among
i nsects the dev elopment o f i ntell igence from instinct as
Whitman would have it o r the independent orig i n s o f i n
Stin c t and i ntell i gence the p robl em o f the i nheritance o f
sychic
acqui
sitions
as
pe
r
Lamarck
the
osition
o
f
natural
p
p
selection as a f actor i n p reservi ng and accumulating favor
able var i ations i n behavior p roblems o f d i stribut i o n etc
Th roughout th i s work the data have given ev i denc e o f
four very den ite att i tudes o f behav i or :
I That there are very den i te and iron clad instinct s
2
That de s pite thes e instincts which are constant i n
each spec i es there i s much variation in the behavior Of
the individuals
I
r
n
x
e
s
s
O
That
there
i
s
a
di
splay
o
f
the
o
f
emo
ions
e
t
p
3
i n these creatures
m
That
in
there
s
uch
aptitude
fo
r
n
i
n
a
n
c
s
i
m
a
s
t
e
4
y
learning d is p lay o f memo ry proting by expe rien ce and
what seems to us i a tio n al conduct
g detai ls S how ing examples o f the above
B e fore g i vi n
four type s o f beha vi o r it i s w ell to po i nt out that the re are
extant two very i m p o rtant i nterpret at ions o f behavi or which
these o b servations clearly do not sup po rt The rst i s the
,
'
'
G E NERAL CO N SI DERATIO N S
35 7
theory o f B ethe w ho thinks that we have not the right to

admit e ither sen s ations p erc eptions or memories i n ani
mals H e re fuses to allow animals any p sychic qualities
whatever and considers the presence o f consciousness
even in animals that can learn by expe rience as a highly
problemati c and improper assumption
The other theory
i s the tropism theory o f Loeb H owever well Loeb s theory
may apply to c ertain organisms i n the p lant or animal
world h is mechan i cal i nterpretation i s n i l in so far as the
movements o f was p s are concerned when he says :

The
trop i sms are i dentical for animals and p lants The ex p la
nati on o f them de p ends rst upo n the S p ecic i rritability
o f certain elements o f the body sur face
and second upon
the relation o f symmetry o f the bod y Symmetric al ele
ments at the sur face o f the b ody h ave the sam e i rritabil ity
unsymmetr i cal elements have a di ffe rent irrita b ility Those
neare r the o ral po le po ssess an i rritabil ity greater than that
o f those near the abo ral pole These c i rcumstances force
an an i mal to o ri ent itsel f i n s uch a w ay that symmetrical
o
i
nts
on
the
su
r
face
o
f
the
body
a
re
stimulated
equally
p
In th i s w ay the an imals are led without w ill o f thei r Ow n
either toward the sourc e o f stimulu s o r away from it Thus
there remains nothi ng for t he ganglion c ells to do but to
conduct the stimulus and thi s may be accompli shed by pro
to p la sm i n any form
2
The theor i es o f Lo eb and B ethe mak e not a beginning of
an e x planation o f the activiti es in Was p do m Had either
o f th e gentlemen s p ent some time i n the eld w ith these
creatures h i s mechanical theo ry i f formulated at all would
,
H o lm es Evolut i o n
,
the
I t m ust b e
reac ti o n s
of
An i m al I n tell igen c e
rem e m b e re d ,
of
an im als
ho
a re
e v e r,
tha t Lo eb
t ro p i s m s
He
r ec o
h av e
a s s o c i ati ve
m em o ry
g iz
n
d o es
b ehav i o r : t rop i s m s , d i ff e ren ti a l s en s ib i li ty an d

H e even s tates that he t hin s he m ust a d m it tha t
20
91
n o t c la im
es
t ha t
al l
three ty p es
of
a s s o c i at iv e
an ts ,
m em o ry
a sp s a n d
b ees
8
35
WA S P S TUD I E S AFI ELD
p resumably have been so modied as not to be all em b rac

i ng in its scope
We know that there are establ ished instincts It would
b e superuous to do more than to ment i on a few o f the
most p atent o f thes e e g the i nstinct o f eve ry S p eci es o f
w a sp to hunt c ertain prey ; the i nstinct for mat i ng ; the i n
s t i n c t for getting food for itsel f ; the i nstinct for stinging
f o r nest
bui ldin g etc These are all establ i shed and funda
mental i nstincts wh ich will adm it o f n o m arked di gression
The re are h o wever an unlim ited numbe r o f habits wh i ch
seem to be constant wh i ch pr o bably are not xed by suc h
stern necess i ty as those ment i on ed above These may b e
mere habits which have b eco m e xed i n the sp ecies even
as our ha b i ts become xed w i thout e i the r the com p uls i on
o f necess i ty o r the u rge o f advantage Nevertheless these
ways seem to be constant w i th i n the spec i es an d seem to b e
followed instinc tively As exam ple s o f th i s condi tion we
may note the follow i ng cas es :
The method o f di gg ing o r o f bui lding i s fai rly constant
amo ng the m em be rs o f a s p e c i es and i s pursued in sti n c
t iv ely
The way o f B em b ix n wbilip en n is in bri nging i n so ft

deli cate ies for the rst meal for h er young i s probably
an act o f i n stin c t at l east i t w ill have to be so class i ed

until otherw i s e proved The large chamber at the bottom
o f the bur row o f th i s spec i es also i s not s o planned by a
knowledge o f the need o f the ba b e nor i s th e work in v ari
ably di rected to thi s end by tropisms but guided again by
inst i nct The constancy w i th which this s p e c l es closes her
hole when goin g out o n a fo raging tr i p c lasses thi s habit
als o w i th instincts The persi stence o f several spec i es o f
wasps in ca r ry i ng the i r prey always in the same way som e
times even going through the moti ons o f drag g in g or tug
g i ng at i t when it i s so smal l that i t i s all the whi le li fted
,
6
0
3
f ound occasional p ronounced variations i n t he type of

nests under our own observation In othe r instances the
nests described by us were di fferent in c ertai n detai l s fr om
those o f the same species described by other wr i te rs f rom
othe r local it i es
P arker tells us that M i c r o b em bi x rst lays the egg and
then provi sions the ne st whil e we found On e which had
brought in v e insect s and st i l l the egg had not been de
pos i ted
A P omp i l oi d es tr op i c us took a S p ider presumably for
her young ; but when s he coul d not nd h er nest and de
cided to abandon i t S he rst made an inci s i on in the body
wal l and sucked the j ui ces
S ome sp ec ies seem to have
quite unvary i ng habits in the manne r o f carrying the i r p rey
but P o mp i l oi des m arg i n a tus exhib i ts s o me degree o f varia
tion i n th is respect ; one walked backwards w i th a sp ider
large r than hers el f holding i t i n a vertical po si tion wi th
her mandibles inserted i n the ventral sur face be tween i ts
legs ; anothe r grasped the spider by one o f i ts coxae ; a
th i rd had a sp i der wh i ch was so small that i t w a s well
above the ground yet she walked backwards w ith it w i th an
ai r o f be i ng obli ged to exert hersel f ; sti ll other s ew back
wards w i th the i r p rey
Wasps are necta r feeding insects yet we nd P tr op ic us
and P m arg i na tus sometimes indulging i n animal fo od
B oth speci es o f P om p i l o i d es st i ng w i th v a ry i n g i ntens i ties ;
sometimes the p rey i s stung to death somet i mes the victim
l i ngers fo r some days or i n othe r i nstances i t recovers
P s c el es tus i ntroduces a novel method o f removi ng the
s oi l from her b urrow by dropp ing he r forelegs t o fo rm a
rake and then walk i ng backwards thus dragging out the
soil which i s usually kicked under the body
P ri oc n em is p o m p i lus i s not a stickl er for conventional ity ;
s he takes such prey as she c a n get uses such hol es as she
.
G ENERAL CO N S I DERATI O N S
1
9
3
can nd an d only sometimes modi e s them to suit her

needs ; i f S he cannot ascend a precipito us S l op e di rectly she
wil l in tim e contrive some ind i rect way to S c a le it I n carry
ing he r p rey S he walks backwards dragging her burden but
also sometimes walks s idew ise dragging i t at right angles to
her own body S he seizes it by a n y o f its members that
may o ffer themselves as a convenient handle at the moment
Other wasps described in the preceding pages p resented
various digress ions f rom the customary w ay o f the species
o f transporting the i r p rey
S ome wasps are very exacting i n their construction or
selection o f a nest ; others admit a considerabl e variety as
for instance Tryp oxy lon c lava tum wh ich i s thus far known
to use fo r i ts cells the m ade over cells o f the mud dauber

holes in logs or fence posts or to tunnel de ftly into wood
to m
ake its o w n burrows
originally a digger in sand
Tac hys p hex term i n a tus
seemed to be adapting itsel f to the l o ose dirt on a clay bank
but the nests we re crude in shape and subnorm al in size
While P ri on onyx thoma e carries the prey to the ori ce
o f the burrow enters come s up grasps it by the antennae
an d pulls i t i n on one occasi on s he hastily dumped it into
the hol e immediately upon arriving and crowded in a fter it
This l ist o f individual digres sions from the customary
ways o f the species might be continued inde nitely
Whethe r these truants from conventional ity S how by thei r
independent actions a type o f degeneration or a deg ree o f
development superior to that o f their fellow beings we
shall not now venture to surm ise ; at least n o t unti l that
moot poi nt has been decided for the human species
However be that as i t may the rst requis ite for improve
ment o f the species or even o f its ability to hol d its own
in a constantly changing envi ronment i s precisely this varia
b ili ty thi s capacity to adopt or contrive new ways
,
2
9
3
Unde r the next heading w e come to the express ion o f

emotions in these wasps I t is exceedingly d i f cult in w it
n ess i n g demonstrations l ike the f ollowing to think o f these
creatu re s as mere automata or m echanical instruments
In the sun dance o f B em b ix n ub i lip en nis su ffi cient emo
tion i s displayed when the dancing males leave thei r aerial
ight and masses o f them struggle on the ground to gain
possession o f the female and then pursue the mated pair
on the wing until they overtake them and form the teeming
mass again
P omp i l oi d es tr op i c us upon regaining her spi der a fter a
long search pounced upon it w ith unmistakable vicio usn es s
Later when some ants came near to he r burrow s he bit at
them kicked at them angrily and curled up her sting at
them menacingly unti l they w ithdrew e ither o f thei r own
accord o r in consequence o f her threats
P omp il oi des n
n a tm
displayed dec i ded agitation
when s he came to a sp ider s turret lingered about it for
some time in va in then v iolently rammed her head against
the tu rret
I f wasps are capable o f the expression o f the emotion
o f anger P s eudag en i a p ttl c hrip en n i s certainly made the
d isplay when the S p i der which she w as pursuing gave one
leap and escaped The wasp b ecame fra ntic ; the s ight o f
the s p i der and the fact that she had been fooled so exc i ted
he r that s he acted madly walking y i ng runn ing abo ut
the ground in indescribable di rections until S he tumbled
headlong i nto a mole s hol e
The play o f Tryp oxy lon
c l a v a tum m ight p robably be classied a s the express ion o f
some emot i on o r other
S ti mts a n i c i nc tm a fter exam i ning near a dozen spots o f
br o ken so i l a rrived at a certain incon sp icuous pile o f loo se
earth Instantly her manner changed when she had struck
pay dirt ; she became greatly excited nervous qu iver i ng

.
W A S P S TU D I E S AFI ELD
9
3 4
o f grass indicated p ractical j udgment to say the least S he

managed it very ingen iously by l i fting on e end o f the bur
den and propp ing i t high in a fork o f grass then swinging
the lowe r end a round to a sl ightly highe r p o int and thus
continued l i fting alternately the ends o f the caterp illar until
she had hoisted i t o ut o f the c has m i n the grass thicket
exactly as a man wo rking alone would ho ist a heavy tim
ber l ittl e by l ittl e
The fact that 5 p i c tip en n is could not be induced to use
the tamp ing tool until the bur row had been nearly lled
shows a marvellou s instinct but the fact that S p hex some
times uses a pebble a ha rd clod a bi t o f w ood o r even the
leg o f a grasshopp er i s evidence o f h ighly plasti c intel
lectual p owe rs When S he throws i n a clod to b e ground
to dust S he breaks i t up ; when i t i s her intention to use
i t as a tool she holds it between the mandibles and p ounds
and rubs it about w ith a c i rcular motion Aga in wh en
s he had le ft her too l in the depression and had so far for
gotten hersel f as to kick loose soi l over i t and bury i t when
w ith he r
s he w a s n ext ready to use i t she went back and
mandibles shed out th is same clod to us e as a hamme r on
the nal laye r
We have wondered as have many others by what sense
the wasp locates the prope r spot for stinging her p rey
We shall not attempt to s ay how S he knows the spot but
the obse rvation o f S p i c tip en n is at thi s crit i cal m oment
shows that the spot i s located by the sense of touch for
S he passed the tip o f he r ab domen on the unde r side o f the
caterpillar feel in g or rathe r probing i n a hal f dozen places
sl iding her sting continuously backward until she reached
the p oint j ust to the rea r o f the last pai r o f p ro legs an d
there admini ste red the long del i berate th rust
The ight o f or ientation as S pi c tip en n is leaves the nest
and the usual return in contrast to the di f culty and s ome
G E NERAL CO N SI DERATI O N S
9
3 5
times the inability when s he leaves the nest or p rey so

suddenly that no study o f the landmarks can be made
aga in demonstrate place memory
Ody n erus g em i n us usually lls up her burro w solid wi th
mud but on one occasion when danger threatened she re
moved enough mud from he r chimney barely to seal the
cell below and then placed a thin layer o f mud ush with
the sur face o f the earth The time w a s too short for the
A l i ttle p i ece o f pape r p inned to the ground to assi st us
i n nding the nest o f O g em i n us proved soon to be a land
mark for this mothe r A fter a few days we removed the
scrap o f pape r ; the mother could not nd the nest until the
paper w a s replaced
Ody n erus d ors al e in going for water follows landmarks
One individual repeatedly ew to the rail road track trav
ell e d parallel t o it fo r a S hort distance then back in the di
rection from which s he came to her p o ol The railroad
track w as undoubtedly associated in her mind w ith the
di rection o f her ight
Fabre lays much stre ss u p on the fact that B embix will
stup idly tolerate dipterous parasites about the nest when
S he ha s the powe r to dispose o f them B ut S tizus un fei ne
fo r she w a s able to recognize at a
tus i s not so S lo w
glance the in festation o f the hopper in the P ri on onyx
hole and ed f rom the place wi thout laying her egg
An Ar ac hn op hoc to n us f errug meus dragg i ng her prey
le ft i t several times to y to the old lo g house some di stance
away and then return ed very di rectly to her sp ider S he
u ndoubtedly travelled from the sp ider to the walls s o o ften
to ascertain i f S he w a s going in the right di rection since
eventually the nest w a s made between its logs
I n P ri oc n emis p omp i lus the instinct to enter the hole each
time be fo re bringing in the sp ide r w a s strong desp ite the
fact that i t meant d i saster to do so fo r the ledge be fore her
,
99
3
doo r w a s s o narrow that every time she came out o f the

burrow she knocked the sp ide r down the hill A fter pro
longed struggles her better j udgment overcame th is in
for s he n ally carried her prey to a
s ti n c t fo r a moment
point where the bank l p ed gradually then dragged i t
back hor i zontally and let i t rest on a ledge three i nches above
the burrow B ut here the conict between instinc t and i n
tellig en c e recurred for when aga in s he tried to take the
spide r i nto her bur row she returned to her old habit o f a
nal i nspection o f the nest and l os t her p rey a s b e fore
Wi th these few facts as examples o f types o f var i at i on
in behavio r plasticity o f in stinct memo ry etc w e can see
at once how little the theory o f B ethe and the tropi sm
theory have to o ffer toward sati s factorily explaining the
3
phenomena Forel conservative as he i s explains the ac
tions o f such c reatures far bette r in these words :
Insects there fore reason and the most intell i gent

among them the social Hym enopte ra especially the wasps
and ants even reason much more than one i s tempted to
beli eve when one observes the regularly recurring mechan
i sm o f the i r i nstincts T o observe and understand these
reasonings well it i s necessary to mi slead thei r instinct
Further one may remark l ittle bursts o f plastic j udgment
.
o f comb i nations extremely l imited i t i s true wh ich in

forc i ng them an instant from the beaten track o f the ir
automati sm helps them to overc o me di fcult i es to decide
,
between two dangers e tc
F rom the point o f v iew o f i nstinct and intell igence or

rather o f reason there are not there fore absolute contrasts
between the insect the mammal and the man In the i n
sect the inher i ted automatisms play by far the p rep o n derat
i ng part develope d and speciali zed to an unheard o f p o int
and curiously enough o ften coming to produce very analo
.
T he S en s es
of
I n s ec t s
T ra n s l a ti o n
1 1
Lo n d o n
9 08
9
8
3
others we real i ze that the student o f an imal b eli av io r must

eve r be on hi s guard against making faulty ej ects ; yet when
con fronted w i th abundant conrmatory evidence to re fuse
'
to ascribe a certa i n psychi c t rait to an animal merely b e

cause it i s not a human be ing seems to us a s great an an
throp o m o rp hi s m as those o f which the older comparative
p sych o logi sts were gu ilty
.
p o m o rp hi c ,
a n im als ,
b ut
t ha t i s to
s ay
also
o th e r
to al l
o c en t r i c
s i s te n t l y
s ho ul
wi f
d ha v e t o
l a ti o n s
a c he
,
k
w hi
ma
we
s ay :
S e n s a t io n s
no
T hi s
f a c i al
e s c e rta i n
c h c o r re s p o n
I t m ay
h ea da c he
c o ul d
t h e re
to m in e i n l i
F
J
o rel
lo n g e r
If
my
s ay
a n im a l
t o th o s e
be
e
at
c o n c ern s
ch
p o s sib le
c a s es,
b ut
t ha t
ren o un c e
as
m a c hi n e
w hi
wi h t
wi f h
whi h I
we
it
hea d a c he
con
We
b eliev e to b e m y
c e rta i n s o n o ro us a rtic u
c
c o n to rti o n s a n d em i t s
f o re
wh
to
o n ly
n ot
I n d ee d b y s up p o s in tha t o ther m en
i tho ut
o u rs el v e s w e a tt ri b u te t o the m
m en
ve the s a m e s e s a tio n s a s
ex ac t p o o f
s ub j e c t i v i s m
our o w
ha
te rm ,
e m it
s he
have
wh
en
m ay
no t a
ha ve
ha v e
r i g ht
to
h ea d
a n a l o g o us
c a ll
them
I N DEX
Ac haeto n e ura 9 2
Ac ri dean n y m p hs 1 48
Ac r os c his m us b ow di c hi
,
2 89
29 4
22 7
hi rs u te , 226-22 7
h ol os eri c ea 2 2 7
-22
2
i
u
l
2
6
i
7
j
20
1
2
1
2
6
e
n
n
i
s
1
i
c
t
i
7
9
,
4
9
p
p
2. gs
2 37 . 29 5 . 36 4-36 5 . g
,
L CKBU
,
33
Am m o bia b ri dw ell i 2 06
Am m op hi la a rg en ta ta 2 27
p es tri s
ca m
B e l o n o g a s t e r, 2
B em b i x c i li a ta 2 7
n u b i li p e n n is , 9 37
62 , 35 8, 360
g s 2 -7
2
i
n
o
l
a
e
I
sp
.
9 . 3. 37 38 . I 47
B E RG S O N 1 , 6
B ETH E. 6, 27 8. 35 7 -35 8. 366
B i cy rtes q ua dif as c ia ta 4 1 -43 g 8
B A
R N a n d CAM E R O N , 1 2 3
B OH N 6
B up r es ti s li ne a ta 1 29
B U I T E L- R E EP E N 2 7 8
C a m p ton o tn s ca r o lin en s is 206
Ca s p i d a e I 3S
Cen th Op hi l us , 2 02
Cera top h o rn s tena x 1 08
Cer c eri s n i tim a 1 33
i
u
m
i
en
n
s
1 29 ,
2
1
6
,
f
p
31
o rna ta, 8
tu b er c ula 2 44 , 2 77
Cha li c o d oma , ex p e ri m en t s
i th
2 78
Cha ly bi on c a er ul eu m 1 2 1 , 1 2 4- 1 2 5 ,
1 35 - 1 37 , 2 88 , g s 30-32
Cha ra c om a n il o ti c a 34 3
C hl o ri o n un rip er 203-206
c a e r ul e um , 44
,
ADAM S 1 7 1
Ag a llia n ovella
Ag en i a a rc hi te c ta 83 84
16
Ag ros ti s C n ig rmn 22 4
Aly s on m ell eus 1 40- 1 44 35 9
,
1 26 - 1 32
rey ,
p a l li d u s
B eetl es
p o li ta 22 7
p roc e ra
,
50
ur na ri a ,
22 7
An thra x
Pa
2 26-2 2 7
5,
2 1 5,
2 26
78
2 4 5 - 2 46
ca p ra ,
68
B DD CDE
37 1 ,
0
34 344 ,
21
n c i s tr o c er us
u lv i p es ,
y a rr o w i ,
Am p h ito rn us ,
t ig ri s ,
2 37 2 43.
s a b ul os a ,
2 26.
215,
rs t
f i r st
n.
m
u
l
o
c
mn
c ol u
8
9
3
others w e real ize that the student o f animal be havior must

eve r be on hi s guard against making faulty ej ects ; yet when
con f ronted w i th abundant conrmatory evidence to re fuse
,
to ascribe a certa i n psychic trait to an an imal merely b e

cause it i s not a human be ing seems to us as great an an
h i m
thro p
a s those o f wh ich the older comparative
psyc h
a n hn z
have
ex a c
s i s te
S h0 1
W lf t
l a ti
'
a ch
s er
I N DEX
0
37
C ra bro
m on ta n n s , 96
o b s c ura s
96
s ex m a cn la tn s , 9 6
Hy p o cra b r o
I S E LE Y
C R E S S ON 9 4
D A VI D S ON 2 02
D A VI S 1 9 8
.
F B
F N A LD 9 7
F li p i ti 38
7
7
97
F li p y 34 9
F L 366 368
1
0- 1
UBB CK
c a esa r,
Ly c i a c og n a ta ria 22 5
Ly c o s a c a r o li n en s i s 5 5
d om if ex 5 8
f r o n di c o la, 4 5 48 ,
hellu o 7 6
Ly c o s i da e, 76 - 79
M C D OU G A LL 1
M a c r os ip hm n rud b ec k i a e
M A R C HA L 8
M A RT I N E A U 2 2 7
M ela n op l us 1 48
d i er en tia l is 1 6 5
1 69
f e m u r rn b m m 1 7 3 1 90
.
G n orim os c h ema g a lla es oli dazg i n i s

336
GROS B E CK , 1 29
G ry llus a b b revia tn s 2 0 1 -2 03
e
n
l
n
s
v
n
a
n
i
c
s
1
p
y
53 1 5 5
Ha li c tus di s p ara lis 1 1 5
p r n i n os n s 1 1 5
s p a rs n s
1 15
v er s a tn s
1 15
HA N O
5 8 1 96
'
C CK
H AN DLI R S CH 2
H A R TMA N 39 60 63
,
24 1 ,
31 9
1 2 8,
1 4 3,
78
309 31
U T ILLI DA
337
A Y
E,
35
Ody n erus a n o rm is 33 1 - 332

d es ig n a tus 332
d ors a li s 62 3 1 2 - 331 365 g 2
g s 5 8
64
r
o
a
m
i
n
t
a
u
s
0
s
f
67
66
334 34
g
e
m
i
n
u
s
2
g
.
9 9 31 2 . 365 . g s 2 . 5 4 .
,
57
44 ,
ed es tri s ,
p
O e d ip o d e
ILLI
2 9 2 . 2 88.
'
GA
H em i p t e ra p rey 1 35
H il a re lla 32 5
Hip p o d a m i a I 3 p un c ta ta 39
HO M E S 2 1 6
H
A RD
H N GER F OR D a n d W
AM S
,
99
32 5 .
N a d a ta g i b b os a 2 40
N EED HA M a n d LLO D 7 7 - 78
N e m o b i n s f as c i a tus 1 5 3
N e o har m o n ia v e lu ti n a 39
N i teli op s i s f os s o r 1 35
i n erm e 1 34
N o tog on zd ea a rg en ta ta
1 5 2- 1 5 8
la ki n n s 1 65
M erm ir ia n e o m ex i c a n a, 1 6 5
M et o p ia , 1 9 7
l eu c o c ep ha la , 1 72
M i c r o b em b i x m on o d o n ta , 39-4 1
-8
0
6
s
3 ,
g
7
M o n o b ia q ua dr i den s 346-35 4
M OR N 2 1 6
M U RTF ELT M i s s M R E , 2 90
1 08
333
G e le c h i i d a e
0- 1 0 1
L
UBB
U
6
1
6
4 47
3 1 64
212
2 36 32 5
c,
2,
re
a ra s
'
ER
21
OR E
es ,
CK
324
LA MA R
35 6
La rra a rg en ta ta, 1 5 6
term i n a ta, 1 49
LATTER, 2 2 7
LEW S 24 5
I OEB 6,
Lo x o s t eg e 306 , 31 0, 336
L
O
2 76 2 7 7
Luc i li a , 9 2
E mp oas ca m a li, 1 43
E n a rm o n i a , 335
E p -i p a sc hi a 349
E um en i d a e 2 , 5 2 99 -35 4
E up a tho c e ra 2 36
E x ertem a 344
A RE 6-8 1 9 1 , 245 277- 2 78
es ,
1
6
32 1
3
D i c er c a p un c tula ta 1 29
D i c hr om orp ha vi ri di s 1 48
D i o d on tus Am eri c a n n s 1 08
c or n i g er, 1 0 7
g r a ci li s , 1 07
m i n u ta s 1 08
tri s tis 1 07 - 1 08
D io m o ru s 94
D i s s os tei ra c a r o li n a 1 6 1 , 1 68 1 7 3
1 75
D o l o m ed e s , 64
D rae c n la c ep ha la m ol lip es 39
E m p ha n op tery x
e um y o thy ro id es
92
s ti rp i c ol us ,
r
r
n
u
i
s
a
s
h
c
g
y
9 4 96
2
1
23
s
0
g
9 94,
'
2
33 334,
1 35
65
I NDEX
Op htha lm o c hln s dn ry i 1 74
O rc h eli m n m d e li ca tm n 1 96
g ra ci l e, 1 9 8
v u lg a r e
1 94, 1 9 6
1 98
O r t ho p te ra , p rey , 1 35 , 1 39 -2 06
O s m i a b ees 27 7
O s p ry n o c ho tn s j un c en s 84
Oxy b e lus q ua dr i n o ta tus 1 0 1
PA
A R D 9 4 1 73 205
P a ra li m n a ap p en di c n la ta 9 8
-1
P a r a n o thy r eus ci n g u la ti s
01
6
9
g s 24 -26
P a rd os a c a na d en s i s 7 6
m i n i m a 76
n ig r ip a lp i s
76
P A R E R 2 2 - 2 4 , 37 , 40 4 1 2 2 7
PE
HA M S 1 9 , 2 3 36-37 , 60-63,
67 . 83. 9 4. 1 08 . 1 1 6- 1 1 9 . 1 2 3.
1 29
1 65 , 2 02 2 2 7 ,
332
P e ll e n es c o ec a tn s 5 7 - 5 8
P el op o eus c a em en ta ri um 1 1 8 - 1 24
1 35 - 1 37 . 2 88 2 5
.
9 . 332 . g s 1 8.
'
om
CK
K
CK
30
32
c o e r ul eus ,
1 1 9 1 20
P e n ta t o m i d a e, 4 2
P ep s i s d u b i ta ta , 67 - 7 1
P hei d o le m n ela n di c a , 39
P h i d ip p us , 60
a n d o x , 84 -8 5
17
,
P hila n thws p un c ta tus , 1 07 - 1 1
7, g s
2, 29
v er ti la b r is ,
1 16
P hl ep s i n s i rr o r at n s , 39
P ho li s o ra ca tull us , 22 4, 3 1 1 ,
324 ,
32 7
P ho rb i a , 9 2
P h or m ia r eg i na 9 2
P ER E . 1 7 4 . 1 9 7 . 2 89 . 2 9 4.
340
P l a n t-li c e, p rey , 1 02 - 1 08
P o l i st e s , 4 - 5 , 286 - 2 88 , g 5 1
a n n u la ri s ,
28 1 2 8 5 , 2 89 - 2 0
9 ,
,
'
I C
b el li c os ns 285 2 9 4
-2
l
li
a
e
s
D
B
D
B
R
2
1
p
p ,
J,
9
94
r u b ig in os us
28 0- 2 83
v a ri a nt s
2 88-2 89 2 9 1
P o m p il i d a e , 4 5 -63
P o m p i l oid es
-6
m a rg i n a tus
8
5
3,
360 362 g 1 2
tr op i c us
45 5 8, 360, 362 , g s
,
- 8
il
u
s
1
p
p
,
7 7
36 1 , 366,
P r i o n on y x a tra tum 24, 62
,
1 75 :
2:
th o m a e, 62
1 7 5 - 1 9 3,
36 1 ,
,
1 1
P o m p i lus s c e les tn s 64, 67 , 36 1

P r i o c n em i s a vi c orn i s 77 -78
,
R ILE Y
R OU B AU D 2
S a li n s p o m p ilus 7 1
S a r c od ex i a s tern o d o n tzs
9 2, 9 4
S a rc o p ha a , 34
S a r c oma c r o n y c hia tri vi tta ta
331
S c a ra b a ei d a e , 1 32
S c eli p hro n s e e P el o o eus
p
S c H WAR z 1 2 4- 1 2 5
S c o l ia d u b ia 1 2 9 - 1 2
3
35 9
S c o lii da e, 2
S
RP
2 26
S i la o n n ig er 1 3 - 1
4 35
S
2 7 8 -2 8o
S N ER 2 9 4
S p a rn op o li us b r evi r os tri s
34 38
S p ha er op h tha lma s c a ev a
89
,
S p he c i us s p e c i os us
44
S p heg i-da e, 2 5
S p he
s e e Am m o hi l a
p
S p hex e l eg a n s 20
3 206
ti b i a li s , 2 0
5
S p i d ers p r ey 6 -6 8 2 7
7 78, 83, 1 1 9
,
'
HA
M I TH
YD
'
P r o d en ia o rn i thog al li , 22
5
P r o mus c a d o m e s ti c a 96
s c e l es tn s ,
64- 67 ,
g 13
P s a m m o c ha ri da e, 5
P s eu da g en ia a r c hi te c ta 83- 8
,
4, g
16
m ellip es , 86 - 89 g s 1 8- 2 0B
p n l c hr ip en n i s 84 -8 5 , 362, g 1 7
P s eu da n thop hi lus ver ti la b ri s 1 1 6
,
1 17
P s i l op us s i p h o, 9 2
P tero m a lus , 84
P y rg us tes s ela ta, 32
4
R a v i n ia q ua dri s et os a
,
91
0
4 42
4
1 59
1
I
37
365 , g s 2 , 4 1
S t rep s i p tera , 3 0
4
S y n a g ri s , 2
S y r ha la a d m i ra b i l i s
,
T a c hy s p hex us c a s
f
.
o b d u c tus
-1
0
5
51
1 48
1 49 - 1
50
I N DEX
2
37
o b s c ura s ,
0
5 15
Vesp a
term i n a tus , 1 44 - 1 49 , 36 1 ,
T e ttig i da e, 1 5 1
34
T e tti g o n i a b id a , 1 43
T ha m m o te tti x c li tel la ri n s ,
Th eri d i n m tep i da r i orum
T H O MA S
T ho m i s i d
73
s p i d e r, 6 2
T hy r eo p us, 96
Tr a c helas tran q ui lla 84
Tro c h os a a va ra , 60,
c i n era
151
c la va tum ,
Try p oxy l on
36 1 . 363. g 32
-I
o
n
h
o
i
1
n
s
j
37 39
trid e n ta tum
1 34
T ry x a l i d, 1 49
,
1 43
1 23
T U R N ER
1 2 4,
225
IL Y
W E S T W OOD
W H EELER 1
1 2.
WIiC K H A M
WILLI AM S
1
1 35 - 1 37 ,
72 ,
1 35 ,
Xy o c eli a
1 49 ,
89
21 5,
51,
5 7,
1 65 ,
2 26
m eta thora c i c us 1 02 - 1 08
g s 27 -2 8
Xy l o c op a vi rg i n i c a 205 346-347
Xy l o m y a p a llip es 9 2
Xy s ti c us n e rv os n s 39
.
78 ,
WILLI S T ON
1 07 .
8 , 1 29
35 6
83
W H I TMA N
L
I
5
m ac n la ta 29 7 -29 8
Ves p u la g erma n i c a 2 9 5 -2 9 7
53
WA S H a n d R E 84
WE S S MA N N 2 78
,
A s u r ve y
an
va l u a t i on
th e
of
ol
der
vi d en c e
d
H e re d
an
f
h
n ew
T
h
e
t
e
o
u
r
n
a
J
l
of
p
I t f urn i s hes th e r s t a uth o r i t a t i v e a c

i ty s a y s :
ork
i
c o un t o f M o r a n s
h
c h i s a va i l a b l e t o a n y
g
o n e b ut t he s e c i a l i s t
A
n d t he
i
!
a
R
l
e
v
e
ew :
p
r e c i a t i on o
an ap
D a r w i n g a ve u s a k n o w l e d ge o f the f a c t o f ev ol u
ti o n , M e n d el i n t r o d u c e d u s t o th e m e tho d o f i n he ri
t a n c e a n d M o rg a n a d v a n c e s u s a l o n g w a y i n u m
d e r s t a n d i n g the m e c ha n i s m
AB O VE
F R E N C H LI N E S
THE
W a l c o tt Am er i c a n a vi a t o r
,
Wa l c o t t
Le t t er s
$
1
ne t ;
f S t u a rt
b y m a il
C h a rl e s D W a l c o tt
S m i th s o n i a n I n s t i t u t i o n w a s a
Dr
S ec r e
t a r y o f t he
c o ll e ge
,
he n h e e n l i s t e d i n t h e F r e n c h F l y i n g
s en i or
C o r p s H e w a s k i l l e d i n c om b a t i n D ec em b e r o f
1917
H i s l ett e r s gi v e on e o f t he b e s t a c c oun ts
e v e r p ub l i s h e d o f th e l i f e o f a n a i rm a n i n t r a i n i n
g
of
s on
E N G LA N D
n a do
An
w a r,
a c c u ra
te
im p
a
ain st
g
WO RLD
ma i l
vol um e
t hi s
ul ty
1 7 4 0- 1 9 1 4
tt e E v er l y S c hm i t t
ev i d e n c e
THE
GE R M A N!
AN D
ar
ti al
P ERIL
P ri n c eto n
$
2
ts
an
y , th e
ne t ;
w i tt
b y m a il
b e f o r e th e
i m p r e s s i ve m a s s o f
e d n a t i on
s e l f outl a
hi s t o ry ,
re s e n
e rm a n
B y B er
en
B y m em b e r s
Un i ve r s it y
of
1
$
t he F a c
n e t;
by
G e rm a n y
m o c ra c y ;
of
vi o l a t i o n
of
Am e ri c a n
i n t e rn a ti o n a l l a w
ri
h
t
s ;
g
f de
I n t er e s t s i n t he w a r
d t he F a r E a s t
G ua r a n
tee s O f i n t e rn a ti o n a l (p e a c e , i n el i m i n a ti o n o f the
P r us s i a n s y s t e m
of
t he t w o A m e r i c a s
an
P R O T E S TA N TIS M I N G E RM AN!
m illan
n e t;
B y K er r D M a c
.
b y m a i l,
d Luthe r a n is m , a n d G e rm a n n a t i on a l
hi c h ha ve m a d e i t p o s s i b l e f or t he m i l
c ha r a c t e r ,
i t a r i s t G e rm a n G o ve rn m e n t t o p l un g e the
o rl d
i n t o t hi s w a r m a d e i n G e r m a n y
Luthe r
an
T H E P R E S I D EN T
TIO NS
the
CO N T R O L
B y E dwa rd S
b y ma i l
A
l et e
s u m m a ry o f
e xec u
t i ve
an
Co r w i n
n et ;
ff a i r s
c o m p r e h en s i v e
By
b y m ai l
ne t ;
l y b e t e en
ov
o f o ur
g
ri va r
d l e g i s l a ti ve b r a n c hes
f o re i gn
in
t he l o n g
CO O P E R A T I VE M A R K E TI N G
la n d
F O REIG N R ELA
c om p
e rn m e n t
OF
s ur v ey
W W
.
Cum b er
of
c o o p er a
ti v e
ro
d uc
d m a rk e t i n g a m o n g the c i t rus f r ui t g r o w e r s
h i c h i ll um i n a t e s the
o f C a l i f o rn i a
hol e el d o f
t i on
'
an
c oop e r a
ti o n
CR I M E P R E VEN T I O N
b y m a il
001
B y Ar th u r
W oo d s
$
1
ne t ;
Wo o d-s
w ho
w as
M a y o r M i t c h el
C om m i ss i on e r t re alt s b ri e y b ut
C o n s t i t u t i o n a l M e t ho d s ;
to p i c s :
,
P ol i c e
t he s e
c o g en t l y
E d uc a t i n g the
P ub l i c ; D i m i n i s h i n g th e S up p l y ; P o v e r t y ; M e n t a l
D e f ec t i ves ; D ri n k a n d D rug s ; C on vi c t s ; J uven i l e
D el i n q u e n c y
.
F I NAN CIN G
50
c en
ts
THE
W AR
B y A B a r t o n H epb u r n
.
b y m a il 5 5
n et ;
c en
ts
M r H ep b u rn s t e xt c ov e r s b ut a s m a l l a r e a o f
r b ut t h a t a r e a i s c ul t i v a t e d i mte n
oo d w h i t e
a
e
p p
g
s i v el y a n d b e a r s a
e n e r o us f rui t a e o f s en s i b l e a n d
g
g
hel p f ul v i e w s o f t he n a t i on s p rob l em i n w a r n a n c e,
b o n d s a n d t a x es
'
S I M P LI F I E D I T ALI A N M A N U A L
t hi n g h a m
5 0 c en t s
net ;
By A L
.
b y m a il 5 5
,
F ro
c en t s .
A l i ttl e b oo k t h a t
i l l b e u se f ul t o a ll w ho
ish
t o l e a r n I t a l i a n q u i c k l y ; es p ec i a l l y , t o !
M C A
m en a n d o the r w a r
o rk e r s
T H E P R OF E S S
S O LD I E R S
b y m a il
A
By
RE
Ldo n
DU C
de
ATI O N
P e a uw
M AIM ED
OF
n et;
l e t e d e s c r i p t i o n o f B el g i um
t o c i vi c u s e f ul n es s th e vi c t i m s
c om p
re s o r e
IO N AL
en
of
d e a vo r
war
to

Wasp Studies

Uploaded by

Document Information

Copyright

Available Formats

Share this document

Share or Embed Document

Sharing Options

Did you find this document useful?

Is this content inappropriate?

Copyright:

Available Formats

Wasp Studies

Uploaded by

Copyright:

Available Formats

3P

Psy cholo gy is an as y e t almost u n explored

a n d se n ses a nd on the o ther up o n tha t

The present volu m e em

A debt o f gratitude i s due to D r C H Turner o f S um

B emb ix n ubi lip en n is

B elonging t o the Fam ily P omp ilida e

P ara n o thy r eus

C era top horus t en ax

Tac hys p hex

A locust hunt er that makes twin cells

Sand loving Ammophila

An ci s troc erus f ulvi p es

Nest o f the p otte r wasp E u me n es s p

The cocoons o f B emb ix n ubi lip en n is

P o mp i l oi d es m a rgi na t us dragging her prey up hill

The remains o f a feast ; cell s o f H yp oc ra b ro sti r

The nest i ng sit e o f P aran o thy r eus c i ng ula ti s

Figure o f female P a ra n o thy reus c i ng ula tis

he r burro w and p rey

Tac hys p hex ter m i na tus

rg ent a ta and he r burrow

C hl o ri on i c hn eum on eum he r bur row and pr ey

The cocoon s o f Sphex p i c tip en n is

wasp s a degree o f i ntelligence which ( w ith the doubt ful

to the human and the illustrious B e rgson using the same

i nsects as paradigms o f instinct i n the sense o f intuiti on

as contrasted w ith intell igence there i s su rely need o f a

WA S P S TUDI E S AFI ELD

per fected nervous and muscular organization a ri d are cer

WA S P S TUDI E S AFI ELD

o f the complete cycl e o f behavior o f any single species can

compared with what can be accompl ished unde r the rigorous

i s commonly prey nest egg and in the E um en id s nest

insti ncts i n the usual b i ological se ns e o f the term They

WA S P S TUDI E S AFI ELD

that they exercise discriminati on and choice that they learn

atomy a v i ew wh ich B ergson i nco rporated i n hi s p hilo s o

present time except among the writers o f certai n nature

Souveni rs w ith del ight frequently ask whether I regard

Origin o f Species appeared to acquire any sympathy w ith

WA S P S TUDI E S AFI ELD

H e i s indeed so p reeminent in the wealth and pre

mentation and in l iterary expression that h i s Souven irs

In one othe r respect al so the Raus have followed a com

Re aumur in hi s M emoi rs publ ished i n 1 734 to 1 74 2 w a s

The B emb ix population burst upon us w ith a suddenness

WA S P STUDI E S AFI ELD

others although sometime s they would conclude w ith one

c ep t ib l e hum o f w ings n o em b rac i n g s no

further domiciles any more than a chick returns to its

WA S P S TUDI E S AFI ELD

e ral minutes or maybe i t w a s seconds So large an assem

reason would immediately po unce upon it w ith a heavy

WA S P S TUDI E S AFI ELD

but there were more o f the vertical emergence holes f rom

WA S P S TUDI E S AFI ELD

cunn i ngly that w e can seldom spy them at thei r work B ut

upon a spot on the bare sur face o f the eld a s p o t w i th

her a comical appearance o f sweeping w ith her elbows

WA S P S TUDI E S AFI ELD

was begun on July 1 4 1 9 1 5 When w e excav ated it on

incomplete o r newly n i shed burrow and we k now that

wh i ch i s one inch long by one hal f inch i n d iameter Thi s