WASPS
SOCIAL AND SOLITARY
[Illustration:
Page 266
PELOPÆUS ON NEST, GROUP OF FINISHED CELLS, AND TUBE OPENED TO SHOW
SPIDERS]
WASPS
SOCIAL AND SOLITARY
BY
GEORGE W. PECKHAM
AND
ELIZABETH G. PECKHAM
WITH AN INTRODUCTION BY
JOHN BURROUGHS
_ILLUSTRATIONS BY JAMES H. EMERTON_
“Bold sons of air and heat, untamed, untired.”—ILIAD, Book XVII
[Illustration: LOGO]
BOSTON AND NEW YORK
HOUGHTON, MIFFLIN AND COMPANY
The Riverside Press, Cambridge
1905
COPYRIGHT 1905 BY GEORGE W. PECKHAM AND ELIZABETH G. PECKHAM
ALL RIGHTS RESERVED
_Published April, 1905_
NOTE
A PART of the matter presented in this volume was published several
years ago by the Wisconsin Biological Survey, under the title
“Instincts and Habits of the Solitary Wasps.” These chapters have
been revised and modified, and new matter based upon later work has
been added, in the hope that in their present less technical form the
observations recorded will be of interest to the general reader.
For a number of the text cuts used in this volume we are indebted to
the courtesy of Dr. E. A. Birge, Director of the Wisconsin Geological
and Natural History Survey.
CONTENTS
CHAPTER PAGE
I. COMMUNAL LIFE 1
II. AMMOPHILA AND HER CATERPILLARS 15
III. THE GREAT GOLDEN DIGGER 56
IV. SEVERAL LITTLE WASPS 72
V. CRABRO 97
VI. AN ISLAND SETTLEMENT 119
VII. THE BURROWERS 141
VIII. THE WOOD–BORERS 178
IX. THE SPIDER–HUNTERS 196
X. THE ENEMIES OF THE GRASSHOPPER 248
XI. WORKERS IN CLAY 265
XII. SENSE OF DIRECTION 275
XIII. INSTINCT AND INTELLIGENCE 292
LIST OF ILLUSTRATIONS
PAGE
PELOPÆUS ON NEST, GROUP OF FINISHED CELLS, AND
TUBE OPENED TO SHOW SPIDERS (page 266) _Frontispiece_
WASP EATING 3
PAPER NEST WITH SIDE REMOVED TO SHOW CONSTRUCTION
OF COMBS 11
AMMOPHILA URNARIA CARRYING CATERPILLAR TO
NEST 19
AMMOPHILA URNARIA STINGING CATERPILLAR 27
CATERPILLAR WITH EGG OF AMMOPHILA URNARIA 29
NEST OF AMMOPHILA 31
AMMOPHILA URNARIA USING STONE TO POUND DOWN
EARTH OVER NEST 39
THOROUGH LOCALITY STUDY BY SPHEX 59
HASTY LOCALITY STUDY BY SPHEX 61
SPHEX DRAGGING GRASSHOPPER TO HER NEST 63
NEST OF SPHEX 69
OXYBELUS QUADRINOTATUS 75
NEST OF OXYBELUS 79
APORUS FASCIATUS 81
WASP HOMES IN THE LOG CABIN 85
NEST OF PERENNIS 89
NEST OF ANORMIS 91
SEXMACULATUS IN THE LINDEN ROOTS 99
CRABRO AND HER WHITE MOTHS 103
CRABRO STIRPICOLA 106
BOTTLE ON STEM TO MEASURE WORK OF CRABRO 107
NEST OF C. STIRPICOLA 113
AMMOPHILA SLEEPING IN THE GRASS (AFTER
BANKS) 115
NEST OF BEMBEX 125
BEMBEX SPINOLÆ LOOKING OUT OF NEST 131
BEMBEX 136
A CORNER OF THE BEMBEX COLONY 137
NEST OF CERCERIS NIGRESCENS 142
CERCERIS CLYPEATA 143
CERCERIS DESERTA: LOCALITY STUDY BEFORE LEAVING
NEST 153
PHILANTHUS PUNCTATUS 157
NEST OF PHILANTHUS PUNCTATUS 163
APHILANTHOPS GATHERING ANTS 169
TRYPOXYLON RUBROCINCTUM 185
MALE TRYPOXYLON AWAITING THE FEMALE 191
TORNADO WASP (POMPILUS QUINQUENOTATUS) DIGGING
NEST 197
POMPILUS QUINQUENOTATUS 199
EPEIRA STRIX PARALYZED AND HUNG UP ON BEAN
PLANT BY POMPILUS QUINQUENOTATUS, OUT OF
THE WAY OF ANTS 203
NEST OF P. QUINQUENOTATUS 213
POMPILUS MARGINATUS 223
THE HOME–COMING OF SCELESTUS 241
NEST OF AGENIA BOMBYCINA 244
LYCOSA KOCHII, FOUND IN NEST OF AGENIA BOMBYCINA
245
TACHYTES 249
NEST OF TACHYTES 251
CHLORION AND THE INDISCREET CRICKET 257
HORIZONTAL CELLS OF THE MUD–DAUBER 271
COURSE FOLLOWED BY POMPILUS FUSCIPENNIS IN
FINDING HER SPIDER, AND IN RETRACING HER
STEPS TO THE NEST 283
LOCALITY STUDY OF ASTATA BICOLOR 288
LOCALITY STUDY OF ASTATA UNICOLOR 289
SECOND LOCALITY STUDY OF ASTATA UNICOLOR 290
PARALYZED SPIDER HUNG UP ON SORREL BY QUINQUENOTATUS
WHILE SHE DIGS HER NEST 295
Introduction
NOT long since I wrote to a friend, a nature lover, as follows: “The
most charming monograph in any department of our natural history that I
have read in many a year is on our solitary wasps, by George W. Peckham
and his wife, of Wisconsin,—a work so delightful and instructive that
it is a great pity it is not published in some popular series of
nature books, where it could reach its fit audience, instead of being
handicapped as a State publication.” This end has now been brought
about, and the book—revised and enlarged with much new material and
many new illustrations—placed within easy reach of all nature lovers,
to whom it gives me pleasure to commend it. It is a wonderful record
of patient, exact, and loving observation, which has all the interest
of a romance. It opens up a world of Lilliput right at our feet,
wherein the little people amuse and delight us with their curious human
foibles and whimsicalities, and surprise us with their intelligence
and individuality. Here I had been saying in print that I looked upon
insects as perfect automata, and all of the same class as nearly alike
as the leaves of the trees or the sands upon the beach. I had not
reckoned with the Peckhams and their solitary wasps. The solitary ways
of these insects seem to bring out their individual traits, and they
differ one from another, more than any other wild creatures known
to me. It has been thought that man is the only tool–using animal,
yet here is one of these wasps, Ammophila, that uses a little pebble
to pound down the earth over her nest. She takes the pebble in her
mandibles, as you or I would take a stone in our hand, and uses it as
a hammer to pound down the soil above the cavity that holds her egg.
This is a remarkable fact; so far as I know there is no other animal on
this continent that makes any mechanical use of an object or substance
foreign to its own body in this way. The act stamps Ammophila as a
tool–using animal.
I am free to confess that I have had more delight in reading this book
than in reading any other nature book in a long time. Such a queer
little people as it reveals to us, so whimsical, so fickle, so fussy,
so forgetful, so wise and yet so foolish, such victims of routine
and yet so individual, with such apparent foresight and yet such
thoughtlessness, finding their way back to the same square inch of
earth in the monotonous expanse of a wide plowed field with unfailing
accuracy, and then at times finishing their cell and sealing it up
without the spider and the egg; hardly any two alike; one nervous
and excitable, another calm and unhurried; one careless in her work,
another neat and thorough; this one suspicious, that one confiding;
one species digging its burrow before it captures its game, others
capturing the game and then digging the hole; one wasp hanging its
spider up in the fork of a weed to keep it away from the ants while
it works at its nest, and then running to it every moment or two to
see that it is safe; another laying the insect on the ground while it
digs,—verily a queer little people, with a lot of wild nature about
them, and of human nature, too.
JOHN BURROUGHS.
WASPS
Social and Solitary
Chapter I
COMMUNAL LIFE
“For where’s the state beneath the firmament
That doth excel the wasps’ for government.”
“What is not good for the swarm is not good for the wasp.”
AS the tendency of mankind to crowd into towns grows stronger the joys
of country life and the workings of Nature are more and more excluded
from the daily experience of humanity. In a few the primal love of
the wild is too strong for suppression, and turning from the hot and
noisy streets they find it a refreshment of spirit to meet our little
brothers of earth and air in the wider spaces of their own territory.
We were walking through the woods one hot day in the middle of August
when our attention was attracted by a stream of yellow–jackets
issuing from the ground. They came in such surprising numbers and
looked so full of energy that we stopped to watch them, and this was
our introduction to the study of these “bold sons of air and heat,”
although a perusal of Fabre’s fascinating “Souvenirs Entomologiques”
had prepared us to feel a lively interest in them. We were at our
summer home near Milwaukee, where meadow and garden, with the wooded
island in the lake close by, offered themselves as hunting grounds,
while wasps of every kind, the socialistic tribes as well as the
extreme individualists of the solitary species, were waiting to be
studied.
The Vespas that had aroused our interest received our first attention,
and a nest in the ground proved to be a most convenient arrangement.
Experiments that would have been dangerous to life and limb had we
tried them with a paper nest hanging in the open, were easy here so
long as we kept calm and unflurried. Intent upon their own affairs,
and unsuspicious of evil, perhaps because they knew themselves to
be armed against aggression, they accepted our presence, at first
with indifference; but as we sat there day after day we must have
become landmarks to them, and perhaps before the summer was over they
considered us really a part of home.
[Illustration: WASP EATING]
While poor humanity takes comfort in a mid–day siesta, wasps love the
heat of noontide, and with every rise in temperature they fly faster,
hum louder, and rejoice more and more in the fullness of life. The
entrance to the Vespa nest was but an inch across; and once when they
were going in and out in a hurrying throng, jostling each other in
their eagerness, we counted the number that passed, one taking the
entrances and one the exits. In ten minutes five hundred and ninety–two
left the nest and two hundred and forty–seven went in, so that we saw
eight hundred and thirty–nine or about eighty to the minute. This must
be a strong swarm, wonderful indeed when we thought that it had all
come from a single queen mother. We imagined how she had made an early
start, digging a hole in the ground, building within it a paper comb
with five or six cells around a central column, and laying therein
some neuter eggs; how she had then spent a month in attending carefully
to the beginnings of things, feeding the young larvæ as they hatched,
and watching over them through their childhood and youth; and then how
her solicitude was rewarded by the filial devotion with which this
first set of workers took upon themselves the labor of excavating,
building, and feeding the young, everything indeed except the
egg–laying. These queens, surrounded though they are by respectful and
attentive subjects, have much the worst of it in our estimation, never
going out, and passing their lives in a dull routine. Through the early
summer only neuters are produced, but when fall approaches the future
generation is provided for by the development of males and females. The
activity of the little colony is limited by the season, for as the days
grow colder the males and females leave the nest and mate, and a little
later both males and workers lose ambition, become inactive and finally
die, while the queens hide away in protected corners to reappear in the
spring. The eggs and larvæ, left unfed and uncared for, become a prey
to moulds and to hordes of insects, and thus the swarm comes to an end.
We had once made some not very successful attempts to find out whether
spiders had a sense of color; and seeing that the conditions were
much more favorable with our present subjects, we thought it would be
a good plan to test their knowledge of the spectrum. Providing six
sheets of stiff paper two feet square, colored respectively red, blue,
green, pink, and two shades of yellow, and cutting a circular hole four
and one half inches in diameter in the centre of each, we began our
experiments by placing the red paper over the nest so that the entrance
was clearly exposed. The outgoing wasps dashed upward without noticing
it, but great was the confusion among the homecomers. Thrown out of
their reckoning, they clamored about us in ever increasing swarms. Like
Homer’s wasps,
“All rise in arms and with a general cry
Assert their domes and buzzing progeny,”
and a crisis (for us) was approaching, when one, a pioneer of thought,
determined to go into the hole, which did not look like the right hole,
although it was where the right hole ought to be; and so potent is
example that one by one the others followed. Three hours later they had
become accustomed to the change, and went in and out as usual.
They had noticed the paper; that was plain enough, but did they notice
the redness? To test this, we left things as they were for two days,
and then substituted blue paper for the red. Again the confusion, the
swarming of fervent legions, the noisy expostulations, the descent of
one after another; but this time they settled down to their ordinary
routine in a little more than two hours. On the following day we
removed the blue paper, leaving the grass around the nest exposed;
and this proved a new source of mystification, but not so serious as
the others. At the end of an hour twenty–five or thirty were still
buzzing about, needing the guidance of the blue paper to get inside,
and entering at once when it was replaced. As we tried new colors
from day to day a few of the wasps became entirely reconciled to our
interference, and paid no attention to the changes, while the others
grew more or less accustomed to the idea of mutability, and were but
little disturbed, although they still showed their consciousness of
each alteration by making a few circles before going in. We once placed
some dark red nasturtiums on light yellow paper near the nest, and
found that more than one third of the homecoming wasps flew to them
and hovered over them before entering. When light yellow nasturtiums,
nearly matching the paper in color, were substituted, only one out of
thirty–six noticed them; and as the odor was as strong in one case as
the other, it would seem that the color was the attracting force.
Our final color experiment was to let the blue paper remain for a day
or two, giving time for all the wasps to become familiar with it, and
then to leave it on the ground a foot and a half away, while replacing
it with yellow. This gave a false nest surrounded by the color that
they had been associating with the entrance, and a true nest surrounded
by a new color. In the next ten minutes two hundred and seventy wasps
came home, and every one of them went to the false nest. Many circled
above it, others entered the hole in the paper, and some began to
excavate, and made quite a depression in the ground; but gradually they
found their way home. Three hours later seventy–six wasps entered the
false nest in five minutes, and at evening they were still visiting
it in goodly numbers; but on the next day we saw only two that were
deceived.
On successive days we substituted red for yellow, green for red, and
so on, always with similar results, although the wasps became more and
more accustomed to the vicissitudes of their life, and after a time
seemed to look for the hole itself without relying upon the color to
guide them. They found their nest under a color new to them much more
readily than when the paper was taken entirely away and the ground
left exposed. Once when the green paper was around their nest, and the
wind blew it over the hole so that they could not enter, at least
one hundred collected, many of them settling in the false nest; when
we lifted the green paper, leaving the hole free, only three or four
entered, but when we put it back in place they rushed in six or seven
at a time. It was plainly the color that directed them.
This was a nearly rainless summer,—a condition extremely favorable
to wasp development. Nests multiplied and grew until the whole
country–side complained, and no wonder, for houses were full of them,
and at mealtimes they gathered at the table with the members of the
family. How did they know when dinner was ready? It could not have been
by the sight, unfamiliar to them, of cooked food; was it, then, through
the sense of smell?
Many were the questions that we asked in vain of our Vespas, but here
was one that they could readily be made to answer. We rolled up two
bundles, one of nothing but gauze, and another, like it in appearance,
but containing some warm chicken bones; these were laid to one side of
the nest, the color of the gauze matching that of the paper on which
it was placed. The wasps in returning to the nest, even though loaded
with food, could not resist the appetizing odor, and settled thickly
upon the bone bundle, trying their best to penetrate within, while
the empty gauze was unnoticed. As the bones grew cold and dry they
attracted less attention, but two days later they were occasionally
visited.
Having killed two wasps that had alighted on the ground, by striking
them with a folded paper, we took them up and placed one of them at a
distance, so that it was entirely hidden in the grass. Five settled
above it, and after they had carried it away the place was visited by
several others, while the spot upon which we had killed them drew to it
nine wasps within fifteen minutes. Thus they seemed very keen of scent
where animal matter was concerned; but the powerful oils of peppermint
and wintergreen, although noticed, aroused little attention, perhaps
because they indicated nothing of interest to them.
Our experiments on hearing met with negative results. The wasps seemed
insensible to any noise we could make or that we could produce by
whistles of various degrees of shrillness. This of course does not show
that they cannot hear, and any one who has been unfortunate enough to
disturb them in the neighborhood of their nest will remember how their
angry buzzing seemed to serve as a battle cry to gather all the members
of the clan for the attack.
Our Vespas began to work an hour or two after sunrise, and did not stop
until dusk. One cloudy evening when darkness fell early they continued
to return to the nest, being able to fly to the right spot without any
hesitation, although our vision did not permit us to see the opening
without going down on our knees and looking closely. At last it grew
perfectly dark, and we stuffed a handkerchief into the hole, with the
result that seventy–five, coming home without a ray of light to guide
them, were shut out, and were found clustered about the spot on the
following morning.
We wanted to estimate the amount of labor done by a worker in a day,
and so, rising one morning at the first bird call, we went out into the
freshness of dawn, and for an hour had the world to ourselves; but a
little before five a few straggling wasps that had stayed out all night
began to bring in loads, and by half past seven they were fairly under
way. From half past four until twelve we counted all that passed, 4534
going out and 3362 coming home; and with all this activity there seemed
to be no pleasure excursions, for each one carried food when returning,
and took out a pellet of earth when leaving. We once raised a little
garden from the pellets that were dropped on our porch table where we
kept a bowl of water. Wasps are great drinkers, and when they find
such a provision they come frequently to refresh themselves, dropping
their loads as they alight. This habit of holding on to their loads
until they settle down may perhaps make them a factor in extending the
boundaries of plant distribution, both under ordinary conditions and
when, as must often happen with little creatures flying so high, they
are blown to long distances from home.
[Illustration: PAPER NEST WITH SIDE REMOVED TO SHOW CONSTRUCTION OF
COMBS]
Having kept close track not only of the numbers, but of the hours,
each count being made to cover five minutes, we were able to calculate
that an average trip occupied forty–three minutes. When we met these
wasps in the garden they never seemed to be hurrying, and had the air
of amusing themselves; but they must be faithful workers to accomplish
so much. The curious fact has been established that when food is very
plentiful the workers begin to lay male eggs, thus taking from the
queen a part of her burden and leaving her free to produce neuters and
females. The nest that we were watching was found, at the end of the
season, to contain 4661 wasps in various stages of development, and
others that we opened had from two to four thousand. This is nothing to
the social wasps of China, where a single household is made up of from
fifteen to twenty thousand members; but China is a thickly populated
country, and perhaps with wasps as with human beings several families
live in a single domicile.
Outside of their wonderful social instincts our wasps are found wanting
in the higher gifts of emotion and intellect. When we killed a number
of them and placed them near the nest, their nearest relatives wasted
no time in mourning, nor yet in revenge, but calmly cut up the bodies
and fed them to the ever hungry young ones. If we placed some rich and
tempting morsels at a distance, two or three would discover them, and
would go back and forth all day without telling the others about it, as
ants would have done under like circumstances. When we obstructed the
opening to their nest by lightly laying blades of grass across, the day
passed without its occurring to the wasps to lift them away, although
they suffered the greatest inconvenience in getting in and out,
crawling laboriously through, and in some instances giving up the task
and flying away.
Vespa maculata, building on trees and fences, has practically the
same habits as the ground wasp, germanica, the internal structure
of the nest following the same plan, while the outer wall is of a
papery substance like that of the combs, made from the scrapings of
weather–beaten wood. The genus Polistes builds combs similar to that of
Vespa, under porches or in any sheltered place, and does not inclose
them. All these wasps, when adult, enjoy fruit and flowers as well
as animal food; but only this last is used for the young, and many a
caterpillar creeping along with sinister design is snatched by them to
be chewed into a pulpy mass, and then fed to the larvæ. No calculation
has been made of the value of these wasps in agriculture, and one
of the things that farmers have yet to learn is to encourage their
presence in orchards and gardens.
Some species are said to sting the drones and larvæ to death at the
close of the season, but this habit is not followed by V. germanica
and V. maculata. Since there is no store of provision to be economized
through the winter the only object of such conduct would be the
merciful one of ending their sufferings at once instead of letting them
perish by slow starvation, and we find no evidence for such elevated
ideas. What makes for the welfare of the species they thoroughly attend
to, but beyond that point they do not go.
The socialism of wasps is in a less evolved state than that of bees and
ants, and yet there is in it sufficient sacrifice of self to the common
good to excite the respectful wonder of human beings, whose relations
to each other and to the state have such different standards.